全 文 ：Journal of Forestry Research, 18(3): 217−220 (2007)

217

DOI: 10.1007/s11676-007-0044-6


Photosynthesis responses to various soil moisture in leaves of
Wisteria sinensis

ZHANG Shu-yong1,2, XIA Jiang-bao3, ZHOU Ze-fu1,2*, ZHANG Guang-can3
1. Research Institute of Forestry, Chinese Academy of Forestry, Beijing, 100091,P, R, China;
2. Key Laboratory of Tree Breeding and Cultivation, State Forestry Administration, Beijing, 100091, P, R, China;
3. Research Center for Eco-Environmental sciences Yellow River Delta, Binzhou University, Binzhou, 256603, Shandong, P, R, China
4. College of Forestry, Shandong Agricultural University, Tai’an, 271018, Shandong, P, R, China

Abstract: A study was conducted to determine the fitting soil moisture for the normal growth of two-year-old W. sinensis (Sims) Sweets by
using gas exchange technique. Remarkable threshold values of net photosynthetic rate (Pn), transpiration rate (Tr) and water use efficiency
(WUE) were observed in the W. sinensis leaves treated by various soil moisture and photosynthetic available radiation (PAR). The fitting
soil moisture for maintaining a high level of Pn and WUE was in range of 15.3%−26.5% of volumetric water content (VWC), of which the
optimal VWC was 23.3%. Under the condition of fitting soil moisture, the light saturation point of leaves occurred at above 800µmol·m-2·s-1,
whereas under the condition of water deficiency (VWC, 11.9% and 8.2%) or oversaturation (VWC, 26.5%), the light saturation point was
below 400µmol·m-2·s-1. Moreover, the light response curves suggested that a special point of PAR occurred with the increase in PAR. This
special point was considered as the turning point that indicated the functional transition from stomatal limitation to non-stomatal limitation.
The turning point was about 600, 1000, 1000 and 400 µmol·m-2·s-1, respectively, at VWC of 28.4%, 15.3%, 11.9% and 8.2% . In conclusion,
W. sinensis had higher adaptive ability to water stress by regulating itself physiological function.
Keywords: Net photosynthetic rate; Soil moisture; Photosynthetic available radiation; Water use efficiency; Wisteria sinensis



Introduction

In recent years, the vegetation restoration and reconstruction
have become an ecological focus, because of the deterioration of
global ecological environment, such as soil and water erosion,
biodiversity loss and global warming etc. Light and soil water
content are important ecological factors influencing plant growth
and distribution (Cai et al. 2004; Cai et al. 2005; Mitton et al.
1998; Zhang et al. 2004). Light has become an outstanding en-
vironmental factor with the decrease of ozonosphere (Huang et
al. 2004; Xu et al. 2005; Zhang et al. 2004). Meanwhile the con-
tradiction between water shortage and demand is becoming acute
in barren mountain area (Cai et al. 2004; Honghton et al. 1996;
Mitton et al. 1998; Xiong et al. 2005; Zeng et al. 2000). There-
fore, how the vegetation adapt to soil drought and light stress,

Foundation project: This research was supported by National Key Sci-
ence and Technology Item in “11th five year” period (No.
2006BAD03A1205), and Shandong Superior Industrial Item in “Breed-
ing and Industrial Exploitation of Superior Liana, Adapting to Afforest-
ing Barren Mountain”.
Received:2007-03-20; Accepted:2007-04-27
©Northeast Forestry University and Springer-Verlag 2007
Electronic supplymentary material is available in the online version of
this article at http://dxdoi.org/10.1007/s11676-007-0044-6
Biography: ZHANG Shu-yong (1980-), male, PH.D., in Research Insti-
tute of Forestry, Chinese Academy of Forestry, Beijing, 100091,P, R,
China. E-mail: zhsyong@126.com
*Corresponding author E-mail: zhouzf@caf.ac.cn
Responsible editor: Hu Yanbo

caused by global climate change, is one of the problems that
people are concerning. Although many studies on the effects of
ecological factors on the ecophysiological characteristics and
space-time dynamics of crops and non-timber forest are currently
available, knowledge of the physiological characteristics espe-
cially photosynthetic responses to soil moisture is scant. By now,
the researches on liana were mainly focused on its development
and utilization (Li et al. 2002; Wu et al. 2004), such as garden
greening, planting technique and officinal value etc., whereas the
study on its ecophysiological characteristics was still at a very
preliminary stage (Huang et al. 2004; Wang et al. 2004), espe-
cially lacking the data of photosynthetic response in leaves of
Wisteriav sinensis (Sims) Sweet to various soil moisture and
light intensity. The aim of this study is to understand the photo-
synthetic characteristics in the leaves of W. sinensis plants under
different soil moisture conditions, which also provides theoreti-
cal support for their use in revegetation of drought regions. Our
objectives are to (1) investigate the effects of soil water deficits
on net photosynthetic rate, transpiration rate and water use effi-
ciency and (2) conform the range of fitting soil water keeping the
leaves higher photosynthetic capacity and water use efficiency.

Materials and methods

General situation of investigation area

The experiment was conducted in forestry experiment station,
which is located in the southeast of Tai’an city (35°38′-36°33′ N,
116°02′-117°59′ E). It belongs to warm temperate zone with a
semi-humid and continental monsoon climate. The average pre-
cipitation for multi-year is 741.8 mm, mainly focused on the 7th
to 9 th month. The annual average temperature is 12.9°C and the
ZHANG Shu-yong et al.

218

warmest month is July. Frostless season is about 202 days.
Brown soil and cinnamon soil are the main soil types.

Materials and water treatment

Eight pieces of two-year-old W. sinensis were used as experi-
mental materials. The plants were potted in March, 2005 (the
pots were buried chronically in the soil to make the same tem-
perature). The soil water conservation and bulk density, meas-
ured by a ring sample, was about 28.1% and 1.17 g/cm3, respec-
tively. Two months later, the plants were irrigated daily to main-
tain 28.4% VWC, and the first measurement was carried out.
Then irrigation was withheld from the third day and the treat-
ment was lasted for 14 days. The change of VWC was monitored
with thetaprobe-MI2X soil moisture probe (AP System, USA).
Photosynthetic parameters of the above leaves were measured at
2 day intervals during the stress period, which were correspond-
ing to VWC 26.5%, 23.3%, 20.9%, 18.4%, 15.3%, 11.9%, 8.2%,
respectively.

Observation of photosynthetic responses to light intensity

Gas exchange measurements were made during the water stress
period, every 2 days between 9:00 and 10:00 h. Every leaf was
studied thrice, and then obtained the mean. Six leaves from each
treatment were used to measure the net photosynthetic rate (Pn),
transpiration rate (Tr), stomatal conductance (Gs) and internal
CO2 concentration (Ci) using a portable photosynthetic system
(CIRAS-2, PP System, UK). The photosynthetic light response
curve was measured at a range of photosynthetic available radia-
tion (PAR), which was changed every 120s in a sequence of
1600, 1400, 1200, 1000, 800, 600, 400, 250, 200, 150, 100, 50
and 20 µmol·m-2·s-1. Leaf-chamber temperature was controlled
under (25±0.5)°C. The following three ratios were calculated:
water use efficiency (WUE) = Pn/Tr (Nijs et al., 1997), stomatal
limitation value (Ls) =1-Ci/Ca (Ca means atmospheric CO2 con-
centration).
Light saturation points (LSP) was calculated by light response
curves of photosynthesis (Pn-PAR curve). Light compensation
points (LCP), dark respiration rate (Rd) and apparent photosyn-
thetic quantum yield (Φ) were calculated by carrying through
linear regression to the beginning parts of response curves
(PAR<200 µmol·m-2·s-1).

Results and analyses

Effect of soil moisture on net photosynthetic rate in leaves of W.
sinensis

The photosynthetic response to light intensity under various soil
moistures was shown as Fig. 1. The leaves became light satu-
rated above 800 µmol·m-2·s-1 in the range of 15.3%-23.3% VWC.
But when VWC was out of this range of soil moisture, leaf LSP
was below 400 µmol·m-2·s-1, of which leaf LSP was at the lowest
value at the point of 8.2% VWC. The change of quantum yield
exhibited a trend similar to photosynthetic rate, which plants at
fitting soil moisture used light more efficiently than those at
lower or higher soil moisture (Fig. 2). According to the Pn-PAR
response curves, remarkable difference in Pn threshold was ob-
served in the leaves treated by various soil moisture. The fitting
soil moisture for the potted W. sinensis was about 15.3%-23.3%.
It was also shown that Φ was about 0.017-0.022, Rd and LCP
was about 0.3-0.8 µmol·m-2·s-1 and 13-51 µmol·m-2·s-1, respec-
tively, when VWC was in the range of 15.3%-26.5%.

Effect of various soil moisture on transpiration rate in leaves of
W. sinensis

As shown in Fig. 3, short-term PAR change had very little influ-
ence on the Tr. Remarkable difference in Tr was observed in
leaves treated by different soil moistures. At water deficiency or
oversaturation, Tr was limited below 1.0 mmol·m-2·s-1. The fit-
ting soil moisture, keeping the leaves a high Tr value was about
20.9%-26.5% VWC.

-1.0
0.0
1.0
2.0
3.0
4.0
5.0
6.0
0 200 400 600 800 1000 1200 1400 1600
Photosynthetic available radiation PAR(µmol·m-2·s-1)
N
et
p
ho
to
sy
nt
he
tic
ra
te
P
n(
µm
ol
·m
-2
·s-
1 )
28.4％ 26.5％ 23.3％
20.9％ 18.4％ 15.3％
11.9％ 8.2％
Fig.1 Light responses of photosynthetic rate
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
0 50 100 150 200
Photosynthetic available radiation PAR(µmol·m-2·s-1)
N
et
p
ho
to
sy
nt
he
tic
ra
te
P
n(
µm
ol
·m
-2
·s-
1 )
28.4％ 26.5％ 23.3％ 20.9％
18.4％ 15.3％ 11.9％ 8.2％
Fig.2 Light responses of photosynthetic rate in low light intensity
0.2
0.6
1.0
1.4
1.8
2.2
2.6
0 200 400 600 800 1000 1200 1400 1600
Photosynthetic available radiation PAR(µmol·m-2·s-1)
Tr
an
sp
ira
tio
n
R
at
e
Tr
(m
m
ol
·m
-2
·s-
1 ) 28.4％ 26.5％ 23.3％
20.9％ 18.4％ 15.3％
11.9％ 8.2％
Fig.3 Light responses of transpiration rate

Journal of Forestry Research, 18(3): 217−220 (2007)

219

Effects of various soil moisture on intercellular CO2 concentra-
tion, stomata limitation and water use efficiency in leaves of W.
sinensis

When VWC was about 18.4%-26.5%, parameters took on a sig-
nificant decrease in Ci and increase in Ls with the increase in
PAR. According to the theory of Farquhar and Sharkey, stomatal
limitation of photosynthesis occurs when Gs and Ci decrease and
Ls increases simultaneously. Therefore it was concluded from
above results that stomatal limitation was the primary factor
influencing Pn in the leaves stressed. However, when VWC was
out of the range (VWC>26.5% or VWC<18.4%), the response
process of Ci and Ls were changed obviously, namely with the
increase in PAR, Ci increased and Ls decreased gradually at ini-
tial stage, whereas PAR was over a special point, Ci decreased
and Ls increased. This special point was considered as the turn-
ing point that indicated the functional transition from stomatal
limitation to non-stomatal limitation. The turning point of PAR
was different under various soil moisture, when VWC was about
28.4%, 15.3%, 11.9% and 8.2%, respectively; the point was
about 600, 1000, 1000 and 400 µmol·m-2·s-1, respectively. It was
inferred from this study that the turning point of PAR was lower
under the unfitting soil moistures than that under the fitting con-
dition.
Leaf WUE was determined by the ratio of Pn and Tr. Thus all
the factors influencing leaf Pn or Tr would change leaf WUE.
When PAR was below 400 µmol·m-2·s-1, WUE increased obvi-
ously with the increase in PAR, otherwise, WUE changed lessly
and maintained a high level in a wide range of light intensity. In
the range of fitting soil moisture (VWC, 15.3%-26.5%), leaf
WUE reached a high level, especially the optimal soil moisture
(23.3%). When soil moisture was higher (VWC, 28.4%) or lower
(VWC, 11.9%), leaf WUE still maintained a relatively higher
level. It was suggested that there was a wide range of soil mois-
ture to maintain the high WUE in leaves of W. sinensis Sweet.

Conclusion and discussion

The process of photosynthesis and transpiration was hypersensi-
tive to the environmental change. The leaf photosynthetic re-
sponse to environment provided the further insights to plant sur-
vival and growth. The Pn, Tr and WUE in leaves of W. sinensis
had the notable threshold value responding to the soil moisture
levels and the variation of PAR. Results showed that the fitting
soil moisture of the leaves was about 15.3%-26.5% (relative
water content (RWC) was about 46.4%-80.3%). This range of
soil moisture not only maintained leaf Pn and WUE at high levels,
but also restrained water consumption arising from transpiration.
When VWC was below 11.9%, severe water stress and high light
intensity usually led to the grievous damage to photosynthesis
apparatus of the plants and the decrease of leaf Pn and WUE (Fig.
1 and Fig. 6). Thus the minimum soil moisture (VWC) was about
11.9% (RWC was about 36.1%), which maintained W. sinensis
normal growth. Previous study showed that not all plants had the
same water requirements and the fitting soil moisture (VWC)
was various among different plants. Usually the fitting soil
moisture in agricultural crops was about 60%-80%, Goldspur
apple tree 60%-71% (Zhang et al. 2004), Black Locust
48%-64%, Oriental Arborvitae 41%-52% ( Zhang et al. 2003;
Zhang et al. 2001). It is inferred from this study that W. sinensis
had strong resistance to drought.

50
100
150
200
250
300
350
0 200 400 600 800 1000 1200 1400 1600
Photosynthetic available radiation PAR(µmol·m-2·s-1)
In
te
rn
al
C
O
2
co
nc
en
tra
tio
n
C
i(µ
m
ol
·m
ol
-1
)
28.4％ 26.5％ 23.3％
20.9％ 18.4％ 15.3％
11.9％ 8.2％
Fig.4 Light responses of internal CO2 concentration

0
0.2
0.4
0.6
0.8
0 200 400 600 800 1000 1200 1400 1600
Photosynthetic available radiation PAR(µmol·m-2·s-1)
St
om
at
a
lim
ita
tio
n
va
lu
e
Ls
(％
)
28.4％ 26.5％ 23.3％
20.9％ 18.4％ 15.3％
11.9％ 8.2％
Fig.5 Light responses of stomata limitation value

-1.0
0.0
1.0
2.0
3.0
4.0
5.0
6.0
0 200 400 600 800 1000 1200 1400 1600
Photosynthetic available radiation PAR(µmol·m-2·s-1)
W
at
er
u
se
e
ffi
ci
en
cy
W
U
E(
µm
ol
·m
m
ol
-1
)
28.4％ 26.5％
23.3％ 20.9％
18 4％ 15 3％
Fig.6 Light responses of water use efficiency

Water stress influenced plant photosynthesis by stomatal
regulation, or directly affected the photosynthetic ability of
mesophyll cells (Castell et al. 1994; Mediavilla et al. 2004;
Tuzet et al. 2003). In the course of water stress, all the respond-
ing process of photosynthesis was divided into three phrases on a
basis of stomatal limitation and non-stomatal limitation (Chen et
al. 2004; Zou et al. 1998). Stomatal limitation mainly involved
locomotion regulation of leaf guard cells, and non-stomatal limi-
tation was correlated with the change of leaf histiocyte (Chen et
al. 2004). The response of Ls and Ci to light intensity indicated
ZHANG Shu-yong et al.

220

that non-stomatal limitation did not occur at the leaves treated by
the fitting soil moisture, but when VWC was over 26.5% or be-
low 18.4%, with the increase in PAR, the factor limited photo-
synthesis was transferred gradually from stomatal limitation
towards non-stomatal limitation, and light use efficiency and
photosynthetic productivity decreased greatly. Results showed
that the turning point of PAR was different in the leaves treated
by various soil moistures (Fig. 4 and Fig. 5).
Plant LCP and LSP reflected the vegetable photosynthetic re-
quirement to light conditions, and embodied its ability to utilize
high light and low light. The vegetation, possessing a low LCP
and high LSP, usually had a strong adaptability to light stress,
contrarily a low adaptability (Ke et al. 2004). It is also shown
that in the range of fitting soil moisture, the LCP of W. sinensis
was about 13-51 µmol·m-2·s-1 (Fig. 2), which was lower than the
limited value of typical heliophiles (50-100µmol.m-2.s-1) (Ke et
al. 2004). It is suggested that W. sinensis can have the shade
tolerance to some extent. The leaves became light saturated
about 800-1000 µmol·m-2·s-1, but W. sinensis still maintained a
high Pn and WUE (Fig. 1 and Fig. 5) in a wider light range
(about 1000-1600 µmol·m-2·s-1).
WUE is not only the comprehensive index for confirming
plant physiological function and adaptability to environment, but
also the initial index ascertaining plant water supply on its de-
velopment and growth (Xiong et al. 2005). When W. sinensis
suffered from water shortage or waterlogging (VWC was over
26.5% or below 15.3%), both Pn and Tr fell down to a low level
under the high light intensity (Fig. 3), but WUE still kept a high
level (Fig. 6).
Based upon all the above discussions, W. sinensis had higher
adaptive ability to water stress by regulating itself physiological
function.

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Chinese Abstracts 3
黑曲霉对二氯甲烷提取物进行抑菌性能测试，结果表明该提
取物对黑曲霉没有抑菌能力。图4表2参29。
关键词:日本花柏；心材外缘；二氯甲烷提取物；气质联机；
抑菌.
CLC number: Q946.8 Document code: A
Article ID: 1007−662X(2007)03-0042-08

07-03-009
ACQ 防腐剂对扭叶松蓝变部分木材力学性能的影响/江京辉
任海青 吕建雄 骆秀琴 吴玉章（中国林业科学研究院木材工
业研究所，国家林业局木材科学与技术重点实验室，北京
100091）//Journal of Forestry Research.-2007, 18(3): 213−216.
本文利用三种不同浓度 ACQ 防腐剂对扭叶松蓝变木材
进行浸注处理，其浓度分别为 1.2%、2.0%和 2.8%。研究其
抗弯弹性模量、抗弯强度、冲击韧性和顺纹剪切强度（弦面）
与未处理蓝变木材相应力学性能的差异，测试标准参照
GB1927~1943-91。研究结果显示，经浸注处理后的试样均达
到美国 AWPA 标准 UC4A 等级规定的药剂保持量；ACQ 防
腐处理大约降低了 20%扭叶松蓝变木材的冲击韧性，与未防
腐处理试样对比，在 0.01水平上有显著差异，但不同浓度间
差异不显著；三种浓度 ACQ处理间以及与未处理的扭叶松蓝
变木材的抗弯强度、抗弯弹性模量和顺纹剪切强度差异不显
著；随着 ACQ浓度的降低，冲击韧性、抗弯强度、抗弯弹性
模量和顺纹弦面剪切强度有所增大，但影响都很小。图 4 表
10 参 6。
关键词：扭叶松，蓝变处理材，非处理材，冲击韧性，抗弯
强度，抗弯弹性模量，顺纹剪切强度（弦面）
CLC number: S781.2 Document code: A
Article ID: 1007−662X(2007)03-0213-04

07-03-010
不同水分条件下紫藤叶片光合作用的光响应/张淑勇(中国林
业科学研究院林业研究所， 北京 100091；国家林业局林木
培育重点实验室，北京 100091)，夏江宝(滨州学院黄河三角
洲生态环境研究中心，滨州 256603)，周泽福(中国林业科学
研究院林业研究所， 北京 100091；国家林业局林木培育重
点实验室，北京 100091)，张光灿(山东农业大学林学院，泰
安 271018） //Journal of Forestry Research.-2007, 18(3):
217−220.
通过测定 2 年生紫藤叶片气体交换参数的光响应，确定
紫藤正常生长发育所需的土壤水分条件。结果表明：紫藤的
光合速率（Pn）、蒸腾速率（Tr）及水分利用效率（WUE）对
土壤湿度和光照强度的变化具有明显的阈值。维持紫藤同时
具有较高 Pn和WUE的土壤湿度范围，在体积含水量（VWC）
为 15.3%～26.5%，其中最佳土壤湿度为 23.3%。适宜的土壤
水分条件下，紫藤光饱和点在 800µmol·m-2·s-1 以上，在水分
不足（VWC为 11.9%，8.2%）或渍水（VWC为 26.5％）时，
光饱和点低于 400µmol·m-2·s-1。此外，光响应曲线表明，随着
光合有效辐射（PAR）增加到一特殊点，气孔限制值（Ls）和
胞间 CO2浓度出现相反的变化趋势。这个点的光合有效辐射
称为光合作用由气孔限制转变为非气孔限制的转折点。并且
不同水分条件下的转折点各不相同，当 VWC 为 28.4%，
15.3% 11.9%和 8.2%，转折点分别为 600， 1000，1000 and 400
µmol.m-2.s-1。总之，紫藤通过对自身生理机能的调节，对水
分胁迫具有较高的适应能力。图 6参 26。
关键词：净光合速率；土壤湿度；光合有效辐射；水分利用
效率；紫藤
CLC number: Q945.11 Document code: A
Article ID: 1007−662X(2007)03-0217-04

07-03-011
传感器数量对应力波检测原木内部缺陷精度的影响/王立海，
徐华东， 周次林， 李莉， 杨学春 (东北林业大学，哈尔滨
150040) //Journal of Foresetry Research.-2007, 18(3): 221−225.
木材无损检测技术是高效利用木材的方法之一。该文阐
述了应力波法检测木材缺陷的原理，分析了传感器数量对图
像的拟合度和误差率两个指标的影响。结果表明，当原木直
径在 20～40cm范围内时，若需对原木缺陷进行精确测量，要
求图像拟合度接近 90%和误差率在 0.1左右时，至少需 12个
传感器才能满足要求；当不需要对原木缺陷进行精确测量，
只需确定缺陷的大致位置时，宜选用 10个传感器进行测量；
当仅仅需要判断原木是否存在缺陷时，选用 6 个传感器就能
满足要求。图 3表 4参 8。
关键词：传感器数量；原木缺陷检测；应力波；图像拟合度
CLC number: S 781.2 Document code: A
Article ID: 1007−662X(2007)03-0221-05

07-03-012
污泥对苗圃生长的银合欢幼苗发芽和初期长势的影
响/G. M. A. Iqbal, S. M. S. Huda*, M. Sujauddin and M. K.
Hossain (Institute of Forestry and Environmental Sciences, Uni-
versity of Chittagong, Chittagong-4331, Bangladesh)//Journal of
Forestry Research.-2007, 18(3): 226−230.
研究了不同类型的污泥（城市的、工业的和住宅污泥）
对苗圃生长的银合欢幼苗田间萌发、生长和分枝的影响。播
种前先将不同类型污泥的混合物与养分匮乏的自然林土壤混
合。播种的 3 和 6 月后，记录幼苗大田发芽、分枝状况和其
他物理生长参数（枝条或根长、活力指数、茎直径、叶片数、
分枝或根鲜重和干重、总的生物量干重增长）等。与对照幼
苗相比，混合污泥的土壤中生长的幼苗田间发芽、分枝状况
及其他生长参数均发生了显著变化。与其它条件生长的幼苗
相比，住宅污泥与土壤混合（1：1）条件生长的 3 月龄和 6
月龄幼苗分枝数和分枝鲜或干重均最高。就生长参数而言，
住宅污泥与自然林土壤混合（1：1）生长的幼苗长势最好。
研究表明：退化的土壤补偿以住宅污泥可促进银合欢的田间
发芽、生长以及分枝的形成。图 1表 3参 29。
关键词：银合欢；污泥；田间发芽；幼苗生长；分枝；活力
指数
CLC number: S718 Document code: A
Article ID: 1007−662X(2007)03-0226-05

07-03-013
鄂尔多斯高原油蒿灌丛群落土壤呼吸日变化和季节变化特征
/金钊(中国科学院地理科学与资源研究所，北京 100101；中