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Review of carbon fixation in bamboo forests in China.

Introduction

Carbon dioxide in the atmosphere has been increasing steadily since 1958. Carbon dioxide (C[O.sub.2]) is one of the so-called 'greenhouse gases' which are responsible for absorbing energy from the sun, leading to warming of the Earth's atmosphere--the 'greenhouse effect. Interest in forest ecosystems has increased due to their role in sequestration and storage of carbon with an increase in greenhouse gas concentration in the atmosphere over recent decades. In forest ecosystems, the carbon storage and the cycling of carbon are common indicators to assess the C[O.sub.2]-fixation capacity. In order to mitigate climate change, more C should be sequestered in forest ecosystems and strategies for an adapted forest management are sought, thus, selection and development of species with a high fixing C[O.sub.2] capacity is of increasing interests worldwide.

Bamboo is a group of plants that belong taxonomically to the subfamily of Bambusoideae under the family of Gramineae and is widespread in the subtropics and tropics of Asia, Africa, and Latin America. There are approximately 1,500 species under 87 genera of bamboo worldwide (Li & Kobayashi, 2004). Bamboo is an important forest type in subtropical and tropical areas. The total area of bamboo forests of various species is 22.0 x [10.sup.6] ha, accounting for about 1.0% of the total area of forest area globally. Although the total forest area has decreased drastically in many countries, bamboo forests have progressively increased at a rate of 3% annually (Guo et al., 2005).

The rate of carbon sequestration depends on the growth characteristics of the tree or bamboo species. Due to their biological characteristics and growth habits, bamboo forests have ecological and environmental functions in soil erosion control, water conservation, land rehabilitation, and carbon sequestration (Zhou et al., 2005). In the global cycles of carbon, bamboo forests are a large carbon sink, but the destruction and degradation of bamboo forests may lead to a carbon source (Li et al., 2003).

In recent years, great advances have been made in fixation carbon ability and carbon storage in the bamboo forest system of China. This paper reviews carbon storage in the vegetation, soil, and litter in bamboo forest systems and compares carbon fixation ability in bamboo forest ecosystems with those of other tree species in subtropical China.

Biological Characteristic and Growth Habits of Bamboo

China is the richest country of bamboo resources in the world with over 500 species in 39 genera and is also a center of the origin and distribution of bamboo. The total area of bamboo forests of various species in China is 7.2 x [l0.sup.6] ha (Zhou et al., 2009).

Phyllostachys pubescen (PP) and Phyllostachys praecox are not only two bamboo species of the greatest planted area in China, but also are those of the greatest economic values and ecological function.

Bamboo is the fastest growing, highest yielding renewable natural resources (Lessard & Chouinard, 1980). Bamboo is known to be one of the fastest growing plants in the world, with a growth rate ranging from 30 to 100 cm/day during the growing season.

In southern China, bamboo is an important forest resource and is also an essential income source for the local farms.

The growth of PP is relatively rapid during the middle of the first month and comparatively slow during the first and last ten days. The time from emergence of the bamboo shoots to formation of young bamboos is only 37-40 d (Zhou & Jiang, 2004). After the formation of the young bamboo, the stem diameters of the bamboo no longer increase, and the increases in height and volume of the stems are also not substantial. Thereafter, storage substance in the bamboo plants increases slowly.

The results of an experiment by Zhou and Jiang (2004) suggested that the growth of bamboo plants is greatly different from the arbors species, and their growth of increasing diameter and height is completed in 40-45 d followed by the slow accumulation of dry matter. The carbon accumulation in bamboo plants are usually completed within one year, and the duration for achieving the greatest carbon storage is much shorter than that of other tree species (>10 years).

Selection cutting is adopted in the management of mature bamboo forests. Usually, 4 or 5-year old bamboo plants are cut down in alternate years. Therefore, PP forests are always in a dynamic equilibrium of growth.

Carbon Storage and its Spatial Distribution in Bamboo Forest Ecosystem

Dynamic Changes in Carbon Element in Single PP Plant and Carbon Accumulation Amounts of One Year-old Bamboo Plants

Zhou and Jiang (2004) reported that the changes in the carbon accumulation in young bamboo plants followed the same pattern as the growth modal. The average C storage of single PP plant ranged from 0.04 kg at the beginning to 1.823 kg at the end of the fast growth period. The average carbon storage (CS) of a single PP plant at the latter period was 44.6 times greater than at the former period. Carbon storage (M) of PP is the function of ground diameter (D), height (H), and the time (t) for emergence of bamboo and is estimated by the following model:

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Study of Zhou et al. (2006) showed that the carbon accumulation amounts of one year-old bamboo plants increased rapidly with time up to 6 months after sprouting of the bamboo shoots and then gradually increased up to 12 months after sprouting. The carbon accumulation amounts by bamboo plants with intensive management (IM) and extensive management (EM) were 10.1 and 5.6 tonnes C/ha, respectively. Carbon accumulation amounts in the old bamboo plants during the first year was very little, accounting for 5.69 to 6.95% of those in the new bamboo ones. The carbon accumulation amount by bamboo plants with IM was 1.8 times as great as with EM. More than 70% of the carbon accumulated in one year-old bamboo plants was contributed by the culms.

Carbon storage (CS) in single plant of PP increased with age classes. However, CS in bamboo stands with class ages of IV (>7- year old bamboo plants), III (5 and 6-year old bamboo plants), II (3 and 4-year old bamboo plants), I (1 and 2-year old bamboo plants) were 21.54, 6.47, 5.32, and 7.55 tC [ha.sup.-1], respectively (Liu et al., 2010).

Spatial Distribution of CS in the Different Organs of Bamboo Plants and CS in Aboveground Parts of Bamboo Stands

Distribution of carbon storage (CS) in different organs of PP was in direct proportion to their biomass. CSs in vegetation story of bamboo stands and its distribution in different organs varied with bamboo species.

Total CS in PP was 30.58 tC/ha, in which the greatest CS was observed in culms (15.587 t/ha), which accounted for 50.97% of total CS, the next greatest CS occurred in roots, and then CS in bamboo stumps, branches, and leaves were lowest (Zhou & Jiang, 2004). Total CS in Pleioblastus amarus (PA) was 46.03 tC/[hm.sup.2], in which the greatest CS was observed in culms (24.426 [tha.sup.-1), which accounted for 53.06% of total CS, the next greatest CS occurred in the leaves and then CS in bamboo rhizomes, branches, bamboo stumps, and roots were lowest (Li et al., 2006). Carbon storage of Neosinocalamus affinis (NA) was 56.27 t/ha, in which the greatest CS was observed in the culms (35.14 t/ha), which accounted for 62.46%, the next greatest CS occurred in the bamboo stumps, and then CS in branches and leaves, and roots were lowest (Wang et al., 2009b).

Carbon accumulation amount in the vegetation of PP stands with IM was 12.75 t/ha/yr, in which carbon accumulation amounts by new bamboo plants and 3-5 year old bamboo plants were 11.39 and 0.648 t/ha/yr, respectively. Carbon accumulation amount in the vegetation of PP stands with EM was 8.144 t/ha/yr, in which carbon accumulation amounts by new bamboo plants and 3-5 year old bamboo plants were 6.056 and 0.421 t/ha/yr, respectively.

Yen and Lee (2011) reported that aboveground CS in bamboo stands were not affected by elevation, but was correlated to stand density (r=-0.600, p=0.039).

Since 1905, bamboo stand area has been increasing year after year, resulting in the increase of CSs in bamboo stands in China. Chen et al. (2009) reported that CSs of bamboo stands in 1950-1962, 1977-1981, 1984-1988, 1989-1993. 1994-1998, and 1999-2003 were 318.55 x [10.sup.12], 427.37 x [10.sup.12], 463.80 x [10.sup.12], 493.00 x [10.sup.12], 548.79 x [10.sup.12], and 631.58 x [10.sup.12] g C, respectively.

Carbon Storage and its Spatial Distribution in Different Bamboo Forest Ecosystems

There are large amounts of carbon in bamboo forest ecosystem and its CS in biomass occupies 22-34% of total CS in bamboo forest ecosystem (Zhou & Jiang, 2004; Li et al., 2006; Wang et al., 2009a).

CSs in bamboo forest ecosystem is related to bamboo species, management, fertilization, etc. Biomass of bamboo forest ecosystems was in the following order: Indosasa crassiflora Mc Olure (182.1 t/ha)>PP (169.4 t/ha)>PA (101.0 t/ha)> Bambusa rigida Keng et Keng f. (83.1 t/ha)>NA(62.3 t/ha)>Dendrocalamus latiflorus Munro (39.2 t/ha) Dendrocalamus latiflorus Munro (10.2 t/ha) (Guo et al., 2005).

Biomass of Indosasa crassiflora Mc Olure under good forest condition (182.1 t/ha) was much greater than that with bad forest condition (126.9 t/ha) (Li et al., 2003). Biomass of high-yield PP forest (169.4 t/ha) was much greater than that of mid-yield one (115.0 t/ha) (Li et al., 2003). CS of PP forest with IM (25.31 t/ha) was greater than that with EM (22.31 t/ha) (Li et al., 2003).

Spatial distribution modal of CS in bamboo forest ecosystem is in the following order: soil layer>plant layer>falling litter>shrub and herb. Total CSs in PP, PA, and NA forest ecosystems were 106.36, 135.81, and 135.95 t/ha, respectively. CS in the soils layer under bamboo forest ecosystems accounted for 67.20, 64.19, and 54.48% of total CS, respectively, CS in the plant layer accounted for 28.75, 33.90, and 21.97% of total CS, respectively; CS in the layer of falling litter, shrub and herb accounted for 4.05, 1.91, and 4.13% of total CS, respectively (Zhou & Jiang, 2004; Li et al., 2006; Wang et al., 2009b).

Carbon Storage in the Litter in Bamboo Forest Ecosystem

In general, CS in the litter in bamboo forest system occupies 2-5% of the total CS. There was a great difference in CS in the litter among different bamboo species. CSs in the litter of bamboo forest ecosystems were in the following order: Indosasa crassiflora Mc Olure (16.6 t/ha)>PP (3.3 t/ha)>Dendrocalamus latiflorus Munro (1.8 t/ha)>Dendrocalamopsis oldhami (Munro) Keng f[approximately equal to]Bambusa rigida Keng et Keng f. (1.2 t/ha) (Li et al., 2003). CS in the litter of Indosasa crassiflora Mc Olure ecosystem is the greatest in the forests studied, which is 7 times greater than that in coniferous and broad-leaf forest ecosystem. CS in the litter of PP forest ecosystem is greater than those in Chinese fir forest, secondary forest, and evergreen broad-leaf forest.

Carbon Storage in the Soils in Bamboo Forest Ecosystem

Generally, CS in forest soils is the greatest carbon stock in terrestrial ecosystem, and its effect occupies an important place in carbon emission and carbon sequestration in terrestrial ecosystem.

Organic carbon in the soils is the greatest carbon stock in bamboo forest system. Organic carbon in the soils usually occupies 54-67% of the total CS. There is a great difference in soil organic contents among different bamboo forest ecosystems. For example, organic content in 0-40 cm of soil depth under PP forest was about 20 g/kg, whereas one under NA forest was up to 30 g/kg (Li et al., 2003).

CSs in the soils under bamboo forest ecosystems depend on organic contents in the soils. CSs in the soil layer of bamboo forests decrease with soil depth. CSs in the soil layer of bamboo forests mainly distributed in 0-20 cm of soil layer, which accounting for about 40% of total CS in 0-60 cm of soil layer (Zhou & Jiang, 2004; Li et al., 2003; Wang et al., 2009a). CS in 0-60 cm of soil layer accounted for 58.2% of total CS in the soil layer, whereas CS in 0-160 cm of soil layer accounted for 89.6% of total CS in the soil layer (Wang et al., 2009a).

The contents of organic matter in the soils under Phyllostachy praecox forests increased with time or cultivation years. The contents of organic matter in the soils in 1999, 2002, and 2007 were 28.9, 44.4 and 55.6 g/kg, respectively. The contents of organic matter in 0-10 cm of soil depth under Phyllostachy praecox forest with 0-, 1-, 5-, 10- and 15-year-cultivation history were 30.96, 25.55, 26.35, 33.07, and 79.24 g/kg, respectively (Jiang et al., 2009).

CSs in the soils under PP forest system ranged 71.48-204.37 t/ha (Zhou & Jiang, 2004; Wang et al., 2009a), whereas CSs in the soils under Phyllostachy praecox forest systems ranged 120.4-271.0 t/ha (Jiang et al., 2009). The soil CSs of the latter were much greater than that the former, which was contributed to heavy-application of organic manure and winter mulching in the management of Phyllostachy praecox production.

According to estimation, CS in soils in bamboo forest system in China was l.066 x [10.sup.15] g C, in which CS in 0-60 cm of soil accounting for 67.20% of total CS.

Total Carbon Storage in Bamboo Forest Ecosystem

Total CSs in bamboo forest ecosystem range 100-243 t/ha. Total CS in bamboo forest systems depends on bamboo species, cultivation ages, mulching, fertilization, soil fertilities, etc.

Total CSs in bamboo forest systems with different bamboo species decreased in the order: NA (135.95 t/ha) (Wang et al., 2009b)[approximately equal to]PA (135.81 t/ha) (Li et al., 2006)> PP (105.7 t/ha) (Liu et al., 2010)>Dendrocalamus latiflorus Munro (10.46 t/ha) (Guo et al., 2005).

Most studies reported that total CSs in the ecosystems of PP stands were greater than that in the ecosystems of Chinese fir and masson pine stands as well as the other forest stands, but few studies showed opposite results (Isagi, 1994; Yen & Lee, 2011; Fan et al., 2006). The reasons for this may be difference in soils, climate, and tree ages. Nonetheless, increase rates of CSs in PP stands are always greater than that in the other forest stands. Isagi (1994) reported that above-ground and belowground net production of bamboo stand was 11.2 and 4.5 tC/ha/year, respectively.

Comparison of CSs in bamboo forests under different management practices suggested that IM reduced total CS by 9.1% as compared with EM (Zhou et al., 2006). The decline in total CS under IM plots contributed to the declines in CS in soil, herb layer, and forest litter.

Total CS in PP forest in Zhejiang Province was estimated by the improved methods (Zhou et al., 2009). The results of calculation showed that total biomass and CS in PP in Zhejiang province were 15.2 x [l0.sup.6] and 7.7 x [l0.sup.6] t, respectively.

The total area of bamboo forests is 4.4 x [l0.sup.6] [hm.sup.2], accounting for 3.6% of the total area of China's forests. According to calculation, carbon storage of vegetation, soils, and litter in bamboo forest ecosystem were 0.2511 x [l0.sup.15], 0.8516 x [l0.sup.15], and 0.036 l x [l0.sup.15] g C, respectively, adding up to 1.1388 x [l0.sup.15] g C (Li et al., 2003).

Comparison of Carbon Fixation Abilities of Different Forest Ecosystems

Bamboo forest ecosystem has a higher potential in fixing C[O.sub.2] from the atmosphere relative to other forest species (Zhou & Jiang, 2004; Zhou et al., 2009; Yuan et al., 2004; Wang et al., 2009a; Yen & Lee, 2011). The annual carbon fixed amount of PP forest ecosystem was greater than the average (7.94 t/ha/a) of forest vegetations in China (Zhao & Li, 1997).Yuan et al. (2004) reported that bamboo forest had the highest carbon fixation ability (14.80 t/ha/a) in 9 kinds of forests studied, with the following order: bamboo forest>slash pine forest> broadleaved forest>rounding trees>cypress forest>masson pine>Chinese fir> open and shrub.

The annual carbon fixation amounts in PP forest ecosystems were 9.9-23.3 tC ha/ (Zhou & Jiang, 2004; Zhou et al., 2006; Zhou et al., 2009; Wang et al., 2009a; Xiao et al., 2007; Yen & Lee, 2011). The annual carbon fixed amount in NA forest ecosystem was 11.25 t/ha/year (Wang et al., 2009b). The annual carbon fixed amounts in PA forest ecosystem was 8.14 t/ha/year (Li et al., 2006).

PP was an uneven-aged stand which was only composed of 1-5-year-old plants, while China fir was an even-aged stand of 15-54 years in tree age, therefore, the mean aboveground carbon sequestration of PP and China fir stands were 8.13 [+ or -] 2.15 and 3.35 [+ or -] 2.02 tC/ha/yr, respectively (Yen & Lee., 2011).

Selective cutting is adopted in the management of mature bamboo stands. Usually, 4 or 5-year old bamboo plants are cut down in alternate years and the thinned bamboo biomass accounts for about one thirds of current total one. Bamboo stands are always in a dynamic equilibrium of growth. Therefore, the mean aboveground carbon sequestration of PP was not dependent on cultivation years. In contrast, the mean aboveground carbon sequestration of Masson pine and China fir stands increase with increasing tree ages up to 12-15 years and decrease slowly thereafter.

Study results of some researchers (Guo et al., 2005; Zhou & Jiang, 2004; Xiao et al., 2007) showed that annual carbon fixation amounts of PP forest ecosystem were much greater than in the ecosystems of Chinese fir and masson pine stands (Ye et al., 2011; Zhou et al., 2009).

Annual carbon fixation amount in NA forest ecosystem were 3.23, 2.21 and 1.38 times greater than Chinese fir (Fang et al., 2002), PP (Zhou & Jiang, 2004), Indosasa crassiflora Mc Olure forest ecosystems (Li et al., 2003), respectively.

Annual carbon fixed amount in the PP forest was 1.39-3.23 and 1.85-2.33 times greater than Chinese fir (Xiao et al., 2007; Zhou et al., 2009; Wang et al., 2009b) and Masson pine stands (Zhou & Jiang, 2004; Zhou et al., 2009), respectively.

IM increased PP numbers per unit areas, which resulted in a great increase in fixation carbon ability. Annual fixation carbon ability of PP forest with IM was 1.56 times as great as with EM (Zhou & Jiang, 2004).

Therefore, it can be concluded that bamboo forest may be one of the favorable forest vegetation for carbon sequestration

Prediction of CS in Bamboo Stand in China and Key Issues to be Studied Further

Based on prediction of development trends of bamboo area in the government strategy documents for forest development over the next five decades, CSs in bamboo stands in 2010, 2020, 2030, 2040, and 2050 are estimated to be 727.08 x [l0.sup.12], 839.16 x [l0.sup.12], 914.43 x [l0.sup.12], 966.803 x [l0.sup.12], and 1017.64 x [l0.sup.12] C, respectively (Chen et al., 2009).

At present, bamboo has been recognized as a species that has the greatest potential in fixing C[O.sub.2] from the atmosphere in subtropics, but it is difficult to determine whether bamboo forest ecosystem is a carbon sink or a carbon source without a study of carbon budget in bamboo forest ecosystem. Furthermore, in the studies of carbon fixation amounts by bamboo forests, a great deal of attention has been focused on PP forest system. Therefore, the key issues to be studied further are as follows:

1. Carbon pools and flux of bamboo forest ecosystems in China;

2. Balance and cycle of carbon in bamboo forest ecosystem;

3. Fate of carbon in bamboo products including bamboo shoots;

4. Exploring the fixation impacts of management including fertilization, mulching, cultivation, etc. on the carbon budget in bamboo forest ecosystem;

5. Exploring the impacts of forest age structure and landuse history on carbon budget in the bamboo forest ecosystem;

6. Soil respiration in the bamboo forest ecosystem;

7. Carbon sequestration ability and carbon budget in PP forest system.

Acknowledgements The authors wish to acknowledge the funding support from the Natural Science Foundation of China (No. 30972356), the Key Science and Technology Development project of Zhejiang Province (2007C 13041) and the Innovative Team Fund (Category B) of Zhejiang Forestry University.

DOI 10.1007/s 12229-011-9082-z

Published online: 14 June 2011

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Guomo Zhou (1) * Cifu Meng (1,2) * Pekun Jiang (1) * Qiufang Xu (1)

(1) Zhejiang Provincial Key Laboratory of Carbon Cycling in Forest Ecosystems and Carbon Sequestration, Zhejiang Agriculture and Forestry University, Lin'an 311300, Zhejiang province, People's Republic of China

(2) Author for Correspondence; e-mail: cifu@21cn.com
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Author:Zhou, Guomo; Meng, Cifu; Jiang, Pekun; Xu, Qiufang
Publication:The Botanical Review
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Date:Sep 1, 2011
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