全 文 ：Journal of Forestry Research, 16(1): 15-18 (2005)

15


Stomatal response of Pinus sylvestriformis to elevated CO2 concentrations
during the four years of exposure

ZHOU Yu-mei, HAN Shi-jie*, LIU Ying, JIA Xia
Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, P. R. China

Abstract: Four-year-old Pinus sylvestriformis were exposed for four growing seasons in open top chambers to ambient CO2 concentration
(approx. 350 µmol·mol-1) and high CO2 concentrations (500 and 700 µmol·mol-1) at Research Station of Changbai Mountain Forest Eco-
systems, Chinese Academy of Sciences at Antu Town, Jilin Province, China (42ºN, 128ºE). Stomatal response to elevated CO2 concentra-
tions was examined by stomatal conductance (gs), ratio of intercellular to ambient CO2 concentration (ci/ca) and stomatal number. Reciprocal
transfer experiments of stomatal conductance showed that stomatal conductance in high-[CO2]-grown plants increased in comparison with
ambient-[CO2]-grown plants when measured at their respective growth CO2 concentration and at the same measurement CO2 concentration
(except a reduction in 700 µmol·mol-1 CO2 grown plants compared with plants on unchambered field when measured at growth CO2 con-
centration and 350 µmol·mol-1CO2). High-[CO2]-grown plants exhibited lower ci/ca ratios than ambient-[CO2]-grown plants when measured
at their respective growth CO2 concentration. However, ci/ca ratios increased for plants grown in high CO2 concentrations compared with
control plants when measured at the same CO2 concentration. There was no significant difference in stomatal number per unit long needle
between elevated and ambient CO2. However, elevated CO2 concentrations reduced the total stomatal number of whole needle by the decline
of stomatal line and changed the allocation pattern of stomata between upper and lower surface of needle.
Key words: ci/ca ratio; High CO2; Pinus sylvestriformis; Stomatal conductance; Stomatal number; Stomatal line
Abbreviations: gs, stomatal conductance; ci, intercellular CO2 concentration; ca, ambient CO2 concentration
CLC number: S718.4 Document Code: A Article ID: 1007-662X(2005)01-0015-04



Introduction

Stomata directly affect the gas exchange of CO2/H2O between
atmosphere and foliage. The impact of elevated CO2 concentra-
tion on the stomatal behavior has attracted considerable attention.
The stomatal response to CO2 is important in understanding
stomatal physiology, and vegetation-atmosphere exchanges at all
scales from the individual plant up to global vegetation (Morison
1998). Short-term and long-term effects of increased CO2 on
stomata are different. In general, the short-term response of sto-
mata is a change in aperture (usually reversible), and long-term
response includes anatomical and morphological changes, for
example, in stomatal number and/or in size (Morison 1998).
Stomatal acclimation may occur when plants are exposed to in-
creased CO2 concentration for a long time.
Stomatal conductance (gs) is most frequently used for assess-
ing the function of stomata in reconciling the water loss and
carbon gain (Zhang et al. 2002). It is generally accepted that an
increase in the ambient CO2 concentration can cause reductions
in stomatal conductance resulted from the decrease of stomatal
aperture and/or density, and the reduction varied widely (Bunce
2000; Morison 2001). A reduction in stomatal conductance is a
common response of herbaceous plants to elevated CO2 (Bunce
2000). Some experimental evidences suggested that many forest
tree species show small or non-significant change in stomatal

Foundation Item: This research was supported by National Basic Research
Program of China (2002CB412502), Project of Key program of the National
Natural Science Foundation of China (90411020) and National Natural Sci-
ence Foundation of China (30400051).
Biography: ZHOU Yu-mei (1973- ), female, Ph. Doctor, assistant research
fellow, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang
110016, P. R. China. E-mail: zhouyumei73@126.com
Received date: 2004-12-01
Responsible editor: Song Funan
*Corresponding author
conductance under long-term elevated CO2 (Curtis 1996; Saxe et
al. 1998), particular conifer (Teskey 1995).
Stomata appear to response directly to the intercellular CO2
concentration (ci), rather than ambient CO2 concentration (ca), as
demonstrated by Mott (Mott 1988). C3 plants normally maintain
relative constancy of the ratio of intercellular to ambient CO2,
approx. 0.7 (Lodge et al. 2001). Given no adjustment of stomata,
the rate of CO2 diffusion through the stomatal pores would rise
in proportion to the increase in ambient CO2 (Jarvis et al. 1999).
Therefore, whether ci/ca ratio remains constant with increased ca
should be examined carefully.
The change of stomatal number of needle is a long-term re-
sponse to elevated CO2 concentration. Some literatures have
reported no change in stomatal density (Poole et al. 2000; Lodge
et al. 2001). There was no difference in stomata density for cur-
rent-year needles of Sitka spruce trees exposed to elevated CO2
concentration for 4 years between treatments (Barton and Jarvis,
1999), the same phenomenon also was observed on Alnus gluti-
nosa (Poole et al. 2000).
The main objectives of this study are to determine stomatal
response of Pinus sylvestriformis to long-term exposure to high
CO2 concentration: (a) to determine gs and ci/ca ratio at different
measurement of CO2 concentration; (b) to examine the changes
of stomatal number of current-year needle.

Materials and methods

The study site is located at Research Station of Changbai
Mountain Forest Ecosystems, Chinese Academy of Sciences at
Antu Town, Jilin Province, China (42ºN, 128ºE). Average annual
rainfall is 700 mm. In 1999, seedlings of Pinus sylvestriformis
were planted in open top chambers and on un-chambered field.
Open top chamber consists of aluminium frames of 1.2 m in
length, 0.9 m in width and height, and clear glass covers. CO2
enters the chamber through perforated plastic pipe at the bottom
ZHOU Yu-mei et al.

16
of chamber. Fan is hung in the top of the chamber to mix the gas
well-proportioned. Treatments consist of three concentration
levels of CO2: ambient, 500 and 700 µmol·mol-1 CO2. Seedlings
in the control chamber and on un-chambered field are given am-
bient CO2, approx. 350 µmol·mol-1. The CO2 concentrations in
each chamber were checked weekly and adjusted. Elevated CO2
concentrations were provided by the mixture of industrial high
CO2 and ambient CO2. 500 and 700 µmol·mol-1 CO2 were ob-
tained by adjusting the velocity and amount of flow of industrial
CO2 and ambient CO2. CO2 concentration was monitored by the
CI—301 gas analyzer once a week. The plants were four years
old, with average height of 47 cm, which are daily irrigated ex-
cept rainy day.
All measurements were made on the current-year needle. The
measured needles were near the top crown and received the full
sunlight. The experiment was begun after the plants had been
exposed to the CO2 treatments for 3 months in the fourth grow-
ing season. Seedlings had been treated by high CO2 concentra-
tions (500 and 700 µmol·mol-1 CO2) continuously (24 h·d-1)
during growing season from June to September since 1999.

Measurement of stomatal conductance and ci/ca ratio
Stomatal conductance (gs) and ci/ca ratio were measured with a
portable photosynthetic analyzer equipped with a conifer cuvette
(LI6400, Li-Cor, Inc., Lincoln, NE). Reciprocal transfer experi-
ment of gs and ci/ca in high- and low-[CO2]-grown plants was
carried out at three levels of CO2 concentrations (350, 500 and
700 µmol·mol-1) in the cuvette, respectively. All measurements
were made directly under light saturating conditions. The read-
ings were taken after allowing gs to reach a steady state.

Stomatal number
Twenty needles were collected at random from 20 plants per
treatment. Stomatal number was separately counted on the upper
and lower surface of current needles. Upper and lower surface of
each needle were cut down 3-mm long epidermis along the nee-
dle, which was viewed with a microscope. The number of all
stomatal lines and number of stomata per line on 3-mm long
epidermis were counted.

Statistics
Mean values of stomatal conductance and ci/ca ratio were
compared separately. One-way analysis of variance was per-
formed for three comparisons. One contrast was carried out to
compare needles grown and measured at 350 µmol·mol-1 CO2
with those grown at 700 and 500µmol·mol-1 CO2 but measured
at 350 µmol·mol-1 CO2. The second contrast was to compare
needles grown and measured at 500 µmol·mol-1 CO2 with those
grown at 700 and 350 µmol·mol-1 CO2 but measured at 500
µmol·mol-1 CO2. The third contrast was to compare needles
grown and measured at 700 µmol·mol-1 CO2 with those grown
at 500 and 350 µmol·mol-1 CO2 but measured at 700
µmol·mol-1 CO2. Stomatal line and stomatal number on upper,
lower surface and whole needle were compared among the four
treatments. All statistical tests were performed using SPSS 11.5
software. The conclusions were reached by the LSD tests.

Results

Reciprocal transfer experiment of gs
Stomatal conductance (gs) of Pinus sylvestriformis in the 500
µmol·mol-1 CO2 was 61% and 4% higher than those in control
chamber and on un-chambered field when measured at their
respective growth CO2 concentrations. Similarly, gs in 700
µmol·mol-1 CO2 increased by 11% and decreased by 28%,
compared with those of growing in the control chamber and on
un-chambered field, respectively (Table 1). The difference was
significant between elevated CO2 and ambient CO2 (p<0.05).
When measured at 500 or 700 µmol·mol-1 CO2, stomatal
conductance of Pinus sylvestriformis at elevated CO2 concentra-
tions was substantially higher than those at ambient CO2 concen-
tration. gs at 700 µmol·mol-1 CO2 was 22% lower than that on
un-chambered field when measured at 350 µmol·mol-1 CO2. gs
at 500 µmol·mol-1 CO2 was the highest at any measurement CO2
concentration. gs of both high-[CO2]-grown and control plants
declined with the increase of measurement CO2 concentration.

Table 1. Mean stomatal conductance (mol·m-2·s-1) in Pinus sylves-
triformis grown at ambient CO2 and elevated CO2 concentrations
measured at three different CO2 concentrations (350, 500, and 700
µmol·mol-1 CO2)
Measurement CO2 concentration
(µmol·mol-1) Growth conditions (µmol·mol-1 CO2)
350 500 700
700 0.284±0.001 0.273±0.0002 0.262±0.0004
500 0.399±0.001 0.379±0.001 0.371±0.002
Control chamber (350) 0.235±0.001 0.216±0.0003 0.206±0.0003
Un-chambered field (350) 0.365±0.004 0.235±0.001 0.181±0.003
Note: Values shown above are means ± standard error. Comparisons
were made among the four treatments at each measurement CO2 concen-
tration. Results for a one-way analysis of variance showed the difference
were significant (p<0.05).

Reciprocal transfer experiment of ci/ca ratio
When measured at their respective growth CO2 concentration,
high-[CO2]-grown plants exhibited lower ci/ca ratios compared
eith the control plants (Table2). ci/ca ratio of Pinus sylvestri-
formis grown at 700 µmol·mol-1 CO2 was 5% and 3% lower
than those at control chamber and un-chambered field, respec-
tively. It was 4% and 2% lower than the control chamber and
un-chambered field for Pinus sylvestriformis grown at 500
µmol·mol-1 CO2. The difference was significant between ele-
vated CO2 and ambient CO2. However, the ci/ca ratio increased
for Pinus sylvestriformis grown in high CO2 concentrations when
exposing to 350 µmol·mol-1 CO2.

Table 2. ci/ca ratio of Pinus sylvestriformis grown at ambient and
elevated CO2 concentrations measured at three different CO2 con-
centrations (350, 500 and 700µmol·mol-1 CO2), respectively
Measurement CO2 concentration
(µmol·mol-1) Growth conditions
(µmol·mol-1 CO2)
350 500 700
700 0.719±0.003 0.685±0.001 0.667±0.001
500 0.728±0.001 0.676±0.001 0.693±0.001
Control chamber (350) 0.701±0.002 0.635±0.002 0.600±0.001
Un-chambered field (350) 0.687±0.002 0.686±0.0003 0.660±0.002
Note: Values shown above are means ± standard error. Comparisons
were made among the four treatments at each measurement CO2 concen-
tration. Results for a one-way analysis of variance showed the difference
were significant except the comparison between 700 µmol·mol-1 CO2
and un-chambered field at 500 µmol·mol-1 CO2 measurement CO2 con-
centration (p<0.05).

Journal of Forestry Research, 16(1): 15-18 (2005)

17
When control plants grown at ambient CO2 concentration were
measured at high CO2 concentrations (500 and 700 µmol·mol-1
CO2) the ci/ca value decreased. The ci/ca ratios of
high-[CO2]-grown plants were higher than those of control plants
when measured at the same CO2 concentration.

Stomatal number
Stomata of Pinus sylvestriformis occur in a few of straight
lines running along the length of the needle on both sides of the
needle. The number of stomatal line per needle (including upper
and lower surface) at high CO2 concentrations was significantly
lower than that on un-chambered field (Table 3). The number of
stomatal line at 700 µmol·mol-1 CO2 approximately equals to
that at 500 µmol·mol-1 CO2. The number of stomatal line of
plants grown on un-chambered field was higher than that in the
control chamber though both accepted ambient CO2 concentra-
tion. The stomatal line of plants in the control chamber was 10%
higher than those at 700 and 500 µmol·mol-1 CO2. But there was
no significant differences between control chamber and elevated
CO2 concentrations. The allocation of stomatal line and number
of stomata was different between upper and lower surface of
needle for high- and low-[CO2]-grown plants. Stomatal line and
number of stomata on upper surface were more than those on
lower surface. The number of stomatal line on the upper surface
of needle grown at 700 and 500 µmol·mol-1 CO2 decreased by
16% and 8%, respectively, compared with that in the control
chamber,. Similarly, the number of stomatal line on the upper
surface of needle grown at 700 and 500 µmol·mol-1 CO2 de-
creased by 25% and 19%, respectively, compared with that on
un-chambered field. The number of stomatal line on the lower
surface of needle grown at 700 µmol·mol-1 CO2 showed no re-
duction and was 11% lower grown at 500 µmol·mol-1 CO2,
compared with that in the control chamber. Pinus sylvestriformis
grown at 700 and 500 µmol·mol-1 CO2 exhibited that the num-
bers of stomatal line on the lower surface of needle were 11%
and 23% separately lower than that at the un-chambered field.
There was no significant difference on the number of stomata on
lower surface among four treatments. Stomatal number per unit
long needle on upper surface at 700 µmol·mol-1 CO2 was much
higher (increased by 16%) than that on the un-chambered field.
However, elevated CO2 did not significantly change the total
stomatal number (including upper and lower surface). The results
of variance on the stomatal line and stomatal number among four
treatments were shown at Table 4.

Table 3. Stomatal line and stomatal number of current-year needle of Pinus sylvestriformis exposure to high CO2 concentrations for four growing
seasons
Growth CO2 concentration (µmol·mol-1 CO2)
Indexes
700 500
Control chamber
(350)
Unchambered field (350)
Number of stomatal line on upper surface of needle 7.6±0.413 8.3±0.442 9.0±0.397 10.2±0.485
Number of stomata 1mm long needle on upper surface 12.1±0.252 11.4±0.249 11.7±0.246 11.3±0.163
Number of stomatal line on lower surface of needle 6.3±0.448 5.5±0.256 6.2±0.296 7.1±0.352
Number of stomata 1mm long needle on lower surface 10. 8±0.171 10.7±0.229 11.0±0.256 10.8±0.183
Number of stomatal line per needle 13.9±0.737 13.8±0.627 15.2±0.601 17.3±0.781
Number of stomata 1mm long needle 22.9±0.359 22.1±0.389 22.6±0.443 22.2±0.294

Table 4. Results of one-way analysis of variance of stomatal line and stomatal number (P: 0.05 level)
1-2 1-3 1-4 2-3 2-4 3-4
Number of stomatal line on upper surface of needle 0.227 0.021 0 0.259 0.003 0.055
Number of stomata 1mm long needle on upper surface 0.056 0.231 0.034 0.464 0.832 0.345
Number of stomatal line on lower surface of needle 0.106 0.838 0.129 0.156 0.002 0.086
Number of stomata 1mm long needle on lower surface 0.693 0.537 0.923 0.315 0.766 0.476
Number of stomatal line per needle 0.959 0.171 0.001 0.156 0.001 0.039
Number of stomata 1mm long needle 0.156 0.591 0.167 0.385 0.971 0.405
Note: 1: 700 µmol·mol-1 CO2; 2: 500 µmol·mol-1 CO2; 3: control chamber; 4: un-chambered field


Discussion

Stomatal conductance, ratio of intercellular to ambient CO2
concentration and stomatal number are main parameters of as-
sessing stomatal behavior at elevated CO2 concentration.
Stomatal conductance can well describe the dynamic changing
trend of stomatal characteristics, while relative stable properties
can be provided by stomatal number (Zhang et al. 2002).
It is widely stated that elevated CO2 concentration will cause
the reduction of stomatal conductance. Stomatal conductance is
affected primarily by stomatal aperture and the number of sto-
mata i.e. stomatal density (Weyers and Lawson 1997). Thus
changes in size and number of stomatal aperture play key roles in
stomatal conductance. Since stomatal number of Pinus sylvestri-
formis was decreased by elevated CO2 concentrations, the in-
crease of stomatal conductance at elevated CO2 concentrations
mainly related to stomatal aperture. Pinus sylvestriformis grown
at 500 µmol·mol-1 CO2 showed the highest stomatal conduc-
tance at any measuring CO2 concentration. However, the change
of stomatal conductance of Pinus sylvestriformis grown at 700
µmol·mol-1 CO2 was related to the measuring CO2 concentration.
Stomata are sensitive to some environmental stimuli, particular
light, humidity and CO2. Therefore, changes of stomatal conduc-
tance in 700 µmol·mol-1 CO2 could be caused mainly by the
change of environmental CO2 concentration. Therefore, the sim-
plest interpretation is that the difference in stomatal conductance
between high- and ambient-[CO2]-grown plants was a result of
the direct adjustment of stomatal aperture.
The change of stomatal aperture or conductance could affect
ZHOU Yu-mei et al.

18
the ci/ca ratio of stomata which are directly sensitive to intercel-
lular CO2 concentration (Mott, 1990). gs and ci/ca ratio in
high-[CO2]-grown plants were higher than those of control plants
when measured at high CO2 concentrations, confirming that
stomata of Pinus sylvestriformis acclimated to long-term expo-
sure to high CO2 concentrations. High-[CO2]-grown plants
showed a decrease for the ratio of ci/ca compared with control
plants when measured at their respective CO2 concentration of
growth. The decrease of ci/ca ratio in high-CO2-grown plants was
mainly caused by higher ambient CO2 concentration (700 and
500 µmol·mol-1 CO2).
Given no adjustment to the change of increasing atmospheric
CO2 concentration the ci/ca ratio would remain constant. How-
ever, Ellsworth (1999) found a similar tendency in ci/ca ratio for
well-watered P. taeda. Sage (1994) and Drake et al. (1997) also
demonstrated it in many experiments, including those in which
plants were grown for long periods in high CO2 concentration.
But it is surprising that the ratio of ci/ca unchanged at high CO2
concentration. Sage (1994) found that except under water and
humidity stress, ci/ca exhibited inconsistent change in a variety of
C3 species. In our study, high-[CO2]-grown Pinus sylvestriformis
exhibited that ci/ca ratio was below the control plants when
measured at their respective growth condition, as also demon-
strated by Wong (1993). ci/ca ratio increased in
high-[CO2]-grown plants when measured under 700 and 500
µmol·mol-1 CO2, confirming that stomata do not always main-
tain ci/ca constant.
Stomatal number of current-year needle of Pinus sylvestri-
formis decreased under 700 and 500 µmol·mol-1 CO2, and the
allocation pattern of stomata between upper and lower surface of
needle gave rise to change. Stomatal lines on the upper surface of
needle showed larger reduction at 700 µmol·mol-1 CO2, and that
on the lower surface showed larger decline at 500 µmol·mol-1
CO2. The difference was not significant though the total stomatal
lines of whole needle (including upper and lower surface) in the
control chamber were higher than that in elevated CO2. The total
stomatal lines of needle on un-chambered field were remarkably
higher than those in the open top chambers (including 700, 500
µmol·mol-1 CO2 and control chamber). Therefore, the microen-
vironment of open top chamber can affect the number of
stomatal line of P. sylvestriformis. That is both elevated CO2 and
microenvironment of open top chamber decreased the number of
stomatal line, but they did not change the stomatal number per
unit long needle. Therefore, the total stomatal number of P. syl-
vestriformis had a decrease at 700 and 500 µmol·mol-1 CO2 by
decreasing stomatal line on upper and lower surface of needle.
The change of stomatal number is a result of long-term exposure
to high CO2 concentrations. The decline of stomatal number for
Pinus sylvestriformis did not significantly affect the change of
stomatal conductance. Some studies showed stomata density did
not change at high CO2 concentration.
There was difference in stomatal response for Pinus sylvestri-
formis to 700 and 500 µmol·mol-1 CO2. The increases of
stomatal conductance and ci/ca ratio of plants grown at 500
µmol·mol-1 CO2 were relatively bigger than those at 700
µmol·mol-1 CO2. In addition, the allocation of stomatal line on
upper and lower surface was different, too. Stomatal behavior of
plants in the control chamber was also different from that on
un-chambered field. By this study, we found that the microenvi-
ronment of open-top chamber affected the physiological charac-
teristics of Pinus sylvestriformis. But we still do not confirm
which factors and how these factors operate.

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Chinese Abstracts 1
《林业研究》（英文）2005年第 16卷第 1期
中文摘要
(Chinese abstracts attached to Journal of Forestry
Research, Vol. 16, No.1 (2005))


05-01-001
利用 Radon 函数变换对木材纹理方向自动检测的研究/于海鹏，刘一星，
刘镇波（东北林业大学生物质材料科学与技术教育部重点实验室, 哈尔滨
150040）//Journal of Forestry Research. –2005, 16(1): 1–4.
本文提出一种利用计算机自动检测木材纹理方向的新方法。四种
Matlab 图像变换函数被尝试用于木材纹理形状的检测。通过比较发现
BWMORPH是最适于检测木材这类中弱纹理的函数，并提取生成了木材
纹理骨骼线图像；再对木材纹理骨骼线图像进行Radon变换，得到 0°~180°
范围内每一角度上的纹理线在投影变换域的积分值，并绘制出纹理线积
分值随角度变化的二维曲线图以反映木材纹理角度上的变化规律。进而
分析了国内 40个树种的纹理方向曲线图以及它们以针叶、阔叶树材和径
向、弦向切面作区别的分类统计规律。结果显示，根据 Radon 变换图及
其纹理曲线图所反映的木材纹理的方向性规律与人们平常对木材纹理的
印象相吻合。这也证明了此种新方法的有效性以及它的应用潜力。图 7
参 6。
关键词：木材；纹理；方向；Radon变换；数字图像处理
CLC Number: S781.3 Document code: A
Article ID: 1007-662X(2005)01-0001-04

05-01-002
基于核糖体 DNA ITS 序列和叶绿体 DNA trnL-F 序列的青篱竹属
Arundinaria 及近缘属系统发育关系的初步研究/诸葛强，丁雨龙，续晨，
邹惠渝，黄敏仁，王明庥（南京林业大学，南京 210037，中国）//Journal
of Forestry Research. –2005, 16(1): 5–8.
在青篱竹属 Arundinaria 系统分类中由于对该属的形态分类标准取
舍不一，至今仍存在着严重的意见分歧。本文采用 PCR扩增产物直接测
序的方法分析青篱竹属 Arundinaria及近缘属（如苦竹属 Pleioblastus、矢
竹属 Pseudosasa、少穗竹属 Oligostachyum、巴山木竹属 Bashania、肿节
竹属 Clavinodum等）有关争议属的代表种或模式种（毛竹为外类群）等
18种竹种的核糖体 DNA ITS区段序列和叶绿体 DNA trnL-F序列。分别
对 ITS序列、 trnL-F序列和 ITS与 trnL-F的组合序列进行系统发育分析。
结果表明，和 trnL-F区段序列相比，核糖体 DNA ITS区段序列有较高的
系统发育信息位点。通过最简约分析产生的 ITS 树、trnL-F 树和 ITS 与
trnL-F 的组合序列树表明，所得树形基本相似。供试竹种（斑苦竹 A.
oleosa、仙居苦竹 A. hsienchuensis、茶秆竹 A. amabilis、长叶苦竹 A. chino、
苦竹 A. amara、宜兴苦竹 A. yixingensis、菲白竹 A. fortunei、翠竹 A.
pygmaea、大明竹 A. gramineus、巴山木竹 A. fargesii、冷箭竹 A. faberi、
凤竹 A. hupehense、鼓节矢竹 P. japonica cv. Tsutsumiana、矢竹 P. japonica、
短穗竹 B. densiflorum、肿节竹 A. oedogonata、少穗竹 A. sulcata）形成一
个单系类群，且分为 2个不同的分支。图 3表 2参 11。
关键词：青篱竹属；核糖体 DNA ITS序列；叶绿体 DNA trnL-F序列; 系
统发育
CLC number: Q344.1 Document code: A
Article ID: 1007-662X(2005)01-0005-04

05-01-003
半干旱黄土高原生物结皮处理下集水造林的初步研究/杨晓晖（中国林业
科学研究院林业研究所，国家林业局林木培育实验室，北京 100091），
王克勤，（西南林学院，昆明 650224），王斌瑞（北京林业大学水土保持
学院，北京 100083），于春堂（中国林业科学研究院林业研究所，国家
林业局林木培育实验室，北京 100091）//Journal of Forestry Research.
–2005-16(1): 9–14.
集水措施是干旱半干旱地区解决因降雨少且季节间分配不均带来的
水资源短缺的主要措施之一，有关微集水区防渗处理材料及其防渗效果
的研究也已广泛开展。本文通过 5 年（1992－1996）的野外试验对半干
旱黄土高原微集水系统生物结皮覆盖对造林的影响进行了研究。结果表
明，结皮接种 3 年后，集水区表面大部分为结皮所覆盖，其集水功能已
经部分地表现出来。在春季 3 个典型时段内浅层土壤含水量与对照相比
显著提高（0.05 水平），1m 土层内春季月均含水量均比对照提高了
0.9%–6.04%。同时在研究期末（1996 年）结皮处理的样地内树木个体的
树高(H)、胸径(DBH)和地径(DGL)均较对照有显著提高（0.05 水平），提
高幅度分别为 22.38%、17.34%和 20.49%。作为一种生物防渗材料，生物
结皮具有无污染、自我繁殖、使用期长和投入小等特点，因此很可以作
为西部大开发中植被恢复的主要集水材料，进一步的工作应放在干旱环
境下当地适宜结皮种类的选择及其快速繁殖技术上。
关键词：造林；生物结皮；微集水区；土壤水分；半干旱黄土高原
CLC number: S728.2 Document code: A
Article ID: 1007-662X(2005)01-0009-06

05-01-004
四个生长季高浓度 CO2 处理长白松的气孔响应/周玉梅，韩士杰，刘颖，
贾夏（中国科学院沈阳应用生态研究所，沈阳 110016）//Journal of Forestry
Research. –2005, 16(1): 15–18.
在长白山站以开顶箱方式对 4 年生长白松连续 4 个生长季进行
CO2处理，包括 700和 500 µmolmol-1高浓度 CO2，以及接受空气 CO2
的对照箱和不扣箱的裸露地条件（约 350 µmolmol-1 CO2），通过测定
气孔导度(gs)，ci/ca比及气孔数量等指标评价气孔对高浓度CO2的响应。
气孔导度及 ci/ca比的转换实验表明，在各自生长 CO2下和在相同测定
CO2下进行比较时，生长在高浓度 CO2下植株的气孔导度要高于空气
CO2下对照组植株的气孔导度（除 700 µmol mol-1 CO2下的植株在生长
CO2浓度下及在 350 µmol mol-1 CO2 下测定时的气孔导度低于裸地植
株外）。在各自生长 CO2浓度下测定时，高浓度 CO2下植株的 ci/ca比
低于对照组植株，但在相同测定 CO2浓度下比较时，却是高浓度 CO2
下植株的 ci/ca高于对照组植株的 ci/ca比。高浓度 CO2下植株与对照组
植株在每单位长度气孔数量上无明显差异，但高浓度 CO2通过降低气
孔线数使长白松当年生针叶的总气孔数量降低，并且改变了气孔在针
叶上、下表面的分配模式。表 4参 18。
关键词：ci/ca 比；高浓度 CO2；长白松；气孔导度；气孔数量；气孔线
CLC number: S718.4 Document code: A
Article ID: 1007-662X(2005)01-0015-04