Volume 50, Issue 3 pp. 265-270

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Growth and Wood/Bark Properties of Abies faxoniana Seedlings as Affected by Elevated CO₂

Yun-Zhou Qiao,

Yun-Zhou Qiao

Chengdu Institute of Biology, the Chinese Academy of Sciences , Chengdu 610041, China

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Yuan-Bin Zhang,

Yuan-Bin Zhang

Chengdu Institute of Biology, the Chinese Academy of Sciences , Chengdu 610041, China

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Kai-Yun Wang,

Corresponding Author

Kai-Yun Wang

Chengdu Institute of Biology, the Chinese Academy of Sciences , Chengdu 610041, China

Shanghai Key Laboratory of Urbanization Processes and Ecological Restoration, East China Normal University , Shanghai 200062, China

*Author for correspondence.
Tel: +86 (0)21 6223 7896;
Fax: +86 (0)21 6223 7896;
E-mail: <[email protected]>.Search for more papers by this author

Qian Wang,

Qian Wang

Chengdu Institute of Biology, the Chinese Academy of Sciences , Chengdu 610041, China

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Qi-Zhuo Tian,

Qi-Zhuo Tian

Agronomic College, Shandong Agricultural University , Taian 271018, China

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Yun-Zhou Qiao,

Yun-Zhou Qiao

Chengdu Institute of Biology, the Chinese Academy of Sciences , Chengdu 610041, China

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Yuan-Bin Zhang,

Yuan-Bin Zhang

Chengdu Institute of Biology, the Chinese Academy of Sciences , Chengdu 610041, China

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Kai-Yun Wang,

Corresponding Author

Kai-Yun Wang

Chengdu Institute of Biology, the Chinese Academy of Sciences , Chengdu 610041, China

Shanghai Key Laboratory of Urbanization Processes and Ecological Restoration, East China Normal University , Shanghai 200062, China

*Author for correspondence.
Tel: +86 (0)21 6223 7896;
Fax: +86 (0)21 6223 7896;
E-mail: <[email protected]>.Search for more papers by this author

Qian Wang,

Qian Wang

Chengdu Institute of Biology, the Chinese Academy of Sciences , Chengdu 610041, China

Search for more papers by this author

Qi-Zhuo Tian,

Qi-Zhuo Tian

Agronomic College, Shandong Agricultural University , Taian 271018, China

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First published: 03 March 2008

https://doi.org/10.1111/j.1744-7909.2007.00604.x

Citations: 3

Supported by the Key Project of the National Natural Science Foundation of China (90511008, 90202010), the Sino-Finland International Cooperative Project (30211130504) and the Program of 100 Distinguished Young Scientists of the Chinese Academy of Sciences (01200108C).

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Abstract

Growth and wood and bark properties of Abies faxoniana seedlings after one year's exposure to elevated CO₂ concentration (ambient + 350 (± 25) μmol/mol) under two planting densities (28 or 84 plants/m²) were investigated in closed-top chambers. Tree height, stem diameter and cross-sectional area, and total biomass were enhanced under elevated CO₂ concentration, and reduced under high planting density. Most traits of stem bark were improved under elevated CO₂ concentration and reduced under high planting density. Stem wood production was significantly increased in volume under elevated CO₂ concentration under both densities, and the stem wood density decreased under elevated CO₂ concentration and increased under high planting density. These results suggest that the response of stem wood and bark to elevated CO₂ concentration is density dependent. This may be of great importance in a future CO₂ enriched world in natural forests where plant density varies considerably. The results also show that the bark/wood ratio in diameter, stem cross-sectional area and dry weight are not proportionally affected by elevated CO₂ concentration under the two contrasting planting densities. This indicates that the response magnitude of stem bark and stem wood to elevated CO₂ concentration are different but their response directions are the same.

Carbon dioxide concentration has been projected to increase from 280 μmol/mol at the beginning of the Industrial Revolution in the middle of the nineteenth century to around 365 μmol/mol today and to continue to rise at about 1.8 μmol·mol⁻¹·y⁻¹ (Mendelsohn and Rosenberg 1994). The effects of elevated CO₂ on plant growth have been studied by ecologists as well as other environmental scientists during the past decades (Conroy et al. 1990; Ceulemans et al. 2002; Kilpeläinen et al. 2005; Handa et al. 2006; Kostiainen et al. 2006).

Tree stems consist of bark and wood. The bark of a tree serves manifold functions, from protective functions to sap conduction of organic materials (Niklas 1999; Du and Yamamoto 2007). In stem wood, vessels function as a transport route for water and various inorganic substances. Fiber provides mechanical support for the stem, and crown and ray parenchyma cells store reserves (Niklas 1999; Du and Yamamoto 2007). Differential effects of elevated CO₂ concentration and plant density may influence plant survival, competition, nutrient and carbohydrate transportation, and timber production (Kaakinen et al. 2004; King et al. 2005). Wood density of some conifers has been shown to increase under elevated CO₂ concentrations (Hättenschwiler et al. 1996; Atwell et al. 2003; Zvereva and Kozlov 2006), but other reports showed no or negative effects of elevated CO₂ (Telewski et al. 1999; Ceulemans et al. 2002; Kilpeläinen et al. 2005). Although there have been some studies on response of plant structure to elevated CO₂, including wood density (King et al. 2005; Kostiainen et al. 2006), wood production (Kilpeläinen et al. 2005) and other characters (Ceulemans et al. 2002; Kaakinen et al. 2004), there are few studies about effects of elevated CO₂ on structural traits of stem bark.

Plant density may vary considerably between different forest stands, including those of monocultures (Wayne et al. 1999). The density can influence tree structure through biomass allocation (Bazzaz et al. 1993), light interception (Bazzaz and McConnaughay 1992), and nutrient transportation (Watt et al. 2005). Studies on density with elevated CO₂ suggested that CO₂-induced growth enhancements were generally lower when individuals were grown in the presence of neighboring plants (Woodward et al. 1991; Retuerto et al. 1996; Wayne et al. 1999). In this study, we investigated the responses of wood and bark characteristics of Abies faxoniana after one growth season's exposure to elevated CO₂ under two planting densities. The aim of the study was to elevate CO₂ concentration to its predicted values for the end of this century (Mendelsohn and Rosenberg 1994). It is hypothesized that stem wood and bark show proportional growth responses to elevated CO₂ concentration.

Results

Elevated CO₂ increased height, stem diameter (D_stem), stem cross sectional area (CSA_stem) and biomass of A. faxoniana seedlings by 9.02 cm, 0.284 cm, 0.426 cm², 3.29 g/plant under low density (elevated CO₂ and low density-ambient CO₂ and low density; ECLD–ACLD) and by 3.33 cm, 0.061cm, 0.064 cm², 1.49 g/plant under high density (elevated CO₂ and high density-ambient CO₂ and high density; ECHD–ACHD, P < 0.05, Table 1). Based on two-way ANOVA (Table 1), CO₂ concentration and plant density both affected all the four parameters shown in Table 1 at significant or highly significant levels.

Table 1. Tree height, stem diameter (D_stem), stem cross-sectional area (CSA_stem), and biomass (mean ± SEM, n = 18) of Abies faxoniana seedlings after exposure for one growing season to elevated CO₂ under different plant density, and summary of ANOVA results

Treatments and source of variance	Growth traits of Abies faxoniana seedlings
Treatments and source of variance	Tree height (cm)		D_stem (cm)		CSA_stem (cm²)		Biomass (g/plant)
ACLD	36.00 ± 2.25a		0.814 ± 0.041a		0.520 ± 0.016a		15.60 ± 1.40a
ECLD	45.02 ± 3.07b		1.098 ± 0.039b		0.946 ± 0.023b		18.89 ± 2.28b
ACHD	28.92 ± 2.58c		0.639 ± 0.047c		0.320 ± 0.037c		8.74 ± 1.24d
ECHD	32.25 ± 2.16a		0.700 ± 0.018a		0.384 ± 0.011c		10.23 ± 1.26c

	SS	P	SS	P	SS	P	SS	P

CO₂	582.54	0.028	0.147	0.000	0.247	0.000	105.67	0.025
Plant density	644.76	0.016	0.390	0.000	0.717	0.000	181.32	0.001
CO₂ × density	485.01	0.031	0.077	0.004	0.065	0.014	33.97	0.393

SS, sum of squares. P, P-value of F-test. Different letters in the same column show difference between treatments significant at P ≤ 0.05, n = 18.

Bark thickness (D_bark), bark cross-sectional area (CSA_bark), bark volume (V_bark) increased with elevated CO₂ but negatively corresponded to density change (Table 2). Bark thickness increased 0.063 cm with elevated CO₂ concentration under the low planting density (P < 0.05) while no effect was found under the high density. Values for CSA_bark and V_bark increased by about 0.173 cm² and 2.91 cm³, respectively, under elevated CO₂ concentration under the low planting density (P < 0.05), and increased by 0.014 cm² and 0.24 cm³, respectively, under the high planting density (P < 0.05). Bark dry weight (DW_bark) was not affected by elevated CO₂ under both densities. Most traits of stem bark were affected by the high planting density, except for D_bark and bark density (DEN_bark) under elevated CO₂. No interactive effect of CO₂ and planting density was identified on bark traits (Table 2).

Table 2. Traits (mean ± SEM, n = 18) of stem bark after exposure for one growing season to elevated CO₂ under different plant density, and summary of ANOVA results

Treatments and source of variance	Stem bark traits of Abies faxoniana seedlings
Treatments and source of variance	D_Bark (cm)		CSA_Bark (cm²)		V_Bark (cm³)		DW_Bark (g bark/plant)		DEN_bark (g DW/cm³)
ACLD	0.103 ± 0.006a		0.106 ± 0.009a		1.27 ± 1.20a		0.37 ± 1.45a		0.29 ± 0.03a
ECLD	0.176 ± 0.005b		0.279 ± 0.019b		4.18 ± 1.94b		0.94 ± 0.04a		0.22 ± 0.04a
ACHD	0.081 ± 0.008a		0.076 ± 0.009c		0.73 ± 0.98c		0.28 ± 0.45b		0.38 ± 0.08b
ECHD	0.088 ± 0.011a		0.090 ± 0.016ac		0.97 ± 1.31ac		0.31 ± 0.80b		0.32 ± 0.06ab

	SS	P	SS	P	SS	P	SS	P	SS	P

CO₂	0.002	0.021	0.011	0.003	7.833	0.001	0.031	0.772	0.010	0.343
Plant density	0.001	0.074	0.015	0.002	10.621	0.000	31.035	0.004	0.013	0.290
CO₂ × density	<0.001	0.325	<0.000	0.551	0.167	0.641	0.549	0.681	<0.000	0.104

Notes are the same as Table 1.

Elevated CO₂ and planting density, either alone or in combination, affected almost all properties of stem wood significantly (Table 3). Wood diameter (D_wood) and wood volume (V_wood) were increased by 0.211 cm and 5.04 cm³, respectively, by elevated CO₂ under the low planting density (P < 0.05), and by 0.054 cm and 0.81 cm³, respectively, under high planting density (P < 0.05). Wood dry weight (DW_wood) showed no reaction, but wood density (DEN_wood) decreased significantly with elevated CO₂ under both planting densities.

Table 3. Traits (mean ± SEM, n = 18) of stem wood after exposure for one growing season to elevated CO₂ under different plant density and summary of ANOVA results

Treatments and source of variance	Stem wood traits of Abies faxoniana seedlings
Treatments and source of variance	D_Wood (cm)		CSA_Wood (cm²)		V_Wood (cm³)		DW_Wood (g wood/plant)		DEN_Wood (g DW/cm³)
ACLD	0.711 ± 0.016a		0.414 ± 0.026a		4.97 ± 0.04a		2.24 ± 0.31a		0.45 ± 0.02a
ECLD	0.922 ± 0.065b		0.667 ± 0.108b		10.01 ± 6.16b		3.83 ± 0.47a		0.38 ± 0.01b
ACHD	0.558 ± 0.039c		0.244 ± 0.031c		2.35 ± 1.25c		1.17 ± 0.86b		0.50 ± 0.04c
ECHD	0.612 ± 0.004a		0.294 ± 0.005a		3.16 ± 2.77d		1.38 ± 0.04b		0.44 ± 0.02a

	SS	P	SS	P	SS	P	SS	P	SS	P

CO₂	0.133	0.000	0.162	0.000	41.741	0.014	0.344	0.825	0.217	0.001
Plant density	0.451	0.000	0.626	0.000	214.624	0.000	49.493	0.001	0.069	0.008
CO₂ × density	0.087	0.000	0.071	0.004	25.930	0.021	2.061	0.593	0.166	0.001

Notes are the same as Table 1.

Influence of elevated CO₂ on bark/wood ratio in diameter, volume and dry weight under the two density levels are shown on Figure 1. Effects of elevated CO₂ differed significantly under the two planting densities. Elevated CO₂ showed significant and positive effects on those variables under the low planting density (P < 0.05) while no effect under the high planting density.

Details are in the caption following the image — **Figure 1**
Open in figure viewer PowerPoint

Bark/wood ratio in diameter (RD), volume (RV) and dry weight (RDW) affected by elevated CO₂ under two planting densities (vertical bars show *SEM* and different letters show difference between treatments significant at P ≤ 0 .05, n = 18).

Discussion

Effects of elevated CO₂

There has been a common prediction that the growth of forest trees in the boreal zone would be enhanced as a result of elevated CO₂, and relevant results have also been reported by many scientists (Woodward et al. 1991; Bazzaz et al. 1993; Yazaki et al. 2004). We find similar trends in our study that tree height, stem diameter, stem CSA as well as biomass increased under conditions of elevated CO₂.

For the stem bark, D_bark, CSA_bark and V_bark increased with elevated CO₂ in our experiment. Yazaki et al. (2004) reported similar a result of increased bark thickness in Japanese Larch under conditions of elevated CO₂ and high nitrogen supplementation. Murthy and Dougherty (1997) reported no change in bark density caused by elevated CO₂. In this study, bark density might be decreased by the fact that V_bark was significantly increased whereas DW_bark was not affected.

Wood volume has been reported to increase under elevated CO₂ in 6-year-old loblolly pine (Telewski et al. 1999) and Scots pine (Ceulemans et al. 2002), but Hättenschwiler et al. (1996) found no stimulation of wood volume in Picea abies in response to elevated CO₂. Atkinson and Taylor (1996) reported a 140% increase in the total stem vessel lumen area caused by the greater vessel number in Quercus robur under conditions of elevated CO₂. Ceulemans et al. (2002), Yazaki et al. (2004) and Kostiainen et al. (2006) also reported increased annual ring width under elevated CO₂. These may provide evidence for stimulated stem wood growth under elevated CO₂ concentrations (Du and Yamamoto 2007; Zhang and Shao 2007). As wood volume is an important criterion of timber productivity, the positive impact of increased CO₂ on wood volume production could be beneficial to silviculture.

Wood density is an important component of wood quality for both timber and paper production (Conroy et al. 1990). A change in DEN_wood may cause a change of wood quality and consequently affect the mechanical and chemical forest industry (Kaakinen et al. 2004; Kostiainen et al. 2004, 2006); for example, the decrease of DEN_wood increased the risk of xylem cavitations (Hacke et al. 2001). DEN_wood has been reported to increase in Pinus radiata due to an elevation of CO₂ concentration from 340 to 660 μmol/mol (Conroy et al. 1990) or remain unchanged (Murthy and Daugherty 1997; Ceulemans et al. 2002). But our results were not in agreement with theirs. Maybe this is caused by the slight enhancement of DW_wood with the larger increase in V_wood. Numerous studies have emphasized the importance of CO₂ enrichment for increasing wood productivity, but many other studies clearly showed that it is associated with change in wood density; for example, plants with high wood density grow slower than those with low wood density (King et al. 2005). The decreased wood density might also mean low wood strength (Kilpeläinen et al. 2005). As changes detrimental for timber strength and commercial use have not been found in current studies (Telewski et al. 1999; Ceulemans et al. 2002; Kostiainen et al. 2004; Kilpeläinen et al. 2005), the speculative offset on the economic benefits of increased productivity by lower timber quality may not necessarily happen under elevated CO₂ environment in future (Watt et al. 2005).

Effects of planting density

Planting density varies considerably in natural forests. However, the reported biomass enhancement of 30–40% resulting from elevated CO₂ is mainly from individual plants raised in the absence of competition (Wayne et al. 1999; Watt et al. 2005). The influence of neighboring plants should be considered in order to improve the predictions of plant growth in response to elevated CO₂ in natural communities and ecosystems (Woodward et al. 1991; Bazzaz and McConnaughay 1992; Körner et al. 1996). It had been hypothesized by Bazzaz et al. (1993) that the magnitude of the growth response to elevated CO₂ is expected to be altered under competitive conditions relative to the responses of plants grown individually. Studies incorporating plant density with elevated CO₂ have drawn the conclusion that CO₂-induced growth was generally lower when individuals were grown in the presence of neighboring plants (Retuerto et al. 1996; Wayne et al. 1999; Watt et al. 2005). Our results agreed well with these researches in that tree height, stem diameter, CSA and biomass of A. faxoniana seedlings were increased by elevated CO₂, and they were lower under high than under low planting density (Table 1).

Such differences between different planting densities could influence future responses of plants to CO₂ enrichment in natural forests where planting density varies over a large range. Contacting and interacting branches and leaves are more abundant with higher planting densities. This may further affect biomass allocation and wood/bark properties in the forthcoming years as it is known that competitive processes modify biomass allocation and wood/bark properties of plants. Thus we can conclude that plant density could play a more important role in response of plants to CO₂ enrichment, as it is usually related to nutrient restriction and stronger competition.

The difference between stem wood and stem bark

In this paper, we hypothesized that stem wood and stem bark of A. faxoniana showed proportional growth with elevated CO₂. However, the proposition is refuted by our observations. The bark/wood ratio in diameter, volume and dry weight were increased in response to elevated CO₂ under the low planting density (Figure 1). This suggests that the response of stem bark to elevated CO₂ was different from stem wood, but their responses were in the same direction because they were both enhanced by elevated CO₂. However, the result can not be extrapolated to mature trees yet, because this was only a short-term experiment on juvenile trees. Experiments in the same region by our working group on more mature trees in natural forest communities are being conducted at present.

In conclusion, this study suggested that growth of A. faxoniana seedlings was stimulated by elevated CO₂ but the response magnitudes were density dependent. Most traits of stem wood and stem bark benefited from elevated CO₂. Nevertheless, effects of elevated CO₂ on wood and bark traits were also density dependent. The result is very important for our understanding of changes in natural forests where planting density varies considerably and competition becomes more complicated than in our mono-species experiments. It also showed that stem bark and wood had similar responses to elevated CO₂, although the magnitudes were different. This suggests that mass allocation between wood and bark changes under enriched CO₂, and in turn, results in variation of wood production and quality. To understand whether these speculations would be true in a future world of elevated CO₂, more attention should be focused on natural forest and mature trees which are of real significance in industrial production.

Materials and Methods

Site descriptions, species and growth conditions

Experiments were carried out in six enclosed-top chambers (ETC) situated in Mao Xian Ecological Research Station (31° 41′ 07″ N, 103° 53′ 58″ E, 1 800 m elevation) which belongs to Chengdu Institute of Biology, Chinese Academy of Sciences. The average CO₂ concentration over one growth season is around 360 μmol·mol⁻¹. Annual precipitation and air temperature were 800 mm and 12 °C, respectively.

Each chamber has a prism as its main body which is made up of a hendecahedron and a corresponding partially full sphere as its top. The horizontal section of the prism is a regular hendecagon (1 m long), and the height of the prism and the sphere are 2 m and 1 m, respectively. They make an internal volume of approximately 24.5 m³, and a ground area of approximately 9.35 m². The side face and top face of the growth chamber are made from 8 mm plate glass (transparency 85%) and 10 mm thick twin-wall-hollow polycarbonate sheet (transparency 82%), respectively. The outside and inside air pressure of the chamber was adjusted by a self-drooping, unilateral pressure-conditioner. A sunshade net, together with the glass wall and polycarbonate top, is used to simulate the real irradiance environment of A. faxoniana seedlings that are grown under natural forest canopy.

A. faxoniana was chosen for study as it is very common in Western China and sometimes occurs as a pioneer species on clear-cutting sites in subalpine coniferous forests (the so-called dark coniferous forests). It is also one of the main constituents of timberline climax communities in the Qinghai-Tibetan Plateau. The seedlings being used were 7 years old. Healthy seedlings of uniform size were chosen and transplanted into experimental wood boxes (60 cm × 70 cm × 40 cm) on 20 September 2003, with a density of 28 or 84 plants/m². Each box was duplicated in each chamber. Boxes were filled with sieved surface sandy soil taken from a 30-year-old natural A. faxoniana forest, and was divided into six portions. Boxes were arranged along the inside glass wall of the chamber. There were two chambers for each CO₂ treatment. The CO₂ concentration was elevated continuously (24 h/day) throughout the growing season from 1 April to 31 September 2004. Seedlings were irrigated to keep the soil water content at 30%–35% according to soil water content logged by a computer, which approximated the optimal water content. Four treatments were classified to test effects of elevated CO₂ and density combined. They are (1) ambient CO₂ and low density (ACLD); (2) ambient CO₂ and high density (ACHD); (3) elevated CO₂ and low density (ECLD) and (4) elevated CO₂ and high density (ECHD).

Measurements and calculations

We measured tree height, diameter, and stem wood diameter of the main stem on 11 September, 2004. Detailed traits measured and methods for measurement are shown in Table 1. Dry weight (DW_t), dry weight of stem bark (DW_bark) and stem wood (DW_wood) are also calculated. Derivative indices of bark and wood traits are calculated as followings:

(1)

(2)

(3)

(4)

(5)

(6)

(7)

Where CSA = cross-sectional area, D = diameter, RD = diameter ratio of bark to wood, RCSA = cross-sectional area ratio of bark to wood, V = volume, DEN_wood = wood density, i = stem or wood. Other variables can be seen in Table 4.

Table 4. Summary of the traits measured on each tree and the methods employed

Trait	Symbols	Units	Method of measurement
Tree height	TH	cm	The distance between the extreme top of main stem and ground.
Stem diameter	D_stem	cm	Means of largest and smallest diameter of the main stem at the ground.
Stem bark thickness	D_bark	cm	Bark is considered to be all tissues external to wood cambium (Hedge et al. 1998; Burrows 2002), which is estimated as the difference of D_stem minus D_wood.
Stem wood diameter	D_wood	cm	Means of largest and smallest diameter of the main stem after peeling.

Statistical analyses

ANOVA was applied to test the effects of elevated CO₂ concentration on traits of stem wood and bark with SPSS-Win 11.5 (SPSS, Chicago, IL, USA). The experimental design is a stochastic one, bust nested, that is, each growth chamber is nested into two CO₂ levels, and two plant densities are nested into chambers. Individual trees were treated as random effects nested within plant density. Individual trees were used as the experimental units thus resulting in a pseudo-replicated design because the variation between trees within chambers was larger in magnitude than the variation between chambers within CO₂ levels (Ceulemans et al. 2002).

(Handling editor: Jian-Xin Sun)

Acknowledgements

The authors thank all the staffs of Mao Xian Ecological Research Station for the donation and help to this experiment.

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Citing Literature

Volume50, Issue3

March 2008

Pages 265-270

Growth and Wood/Bark Properties of Abies faxoniana Seedlings as Affected by Elevated CO₂

Abstract

Results