全 文 ：第 23 卷　第 3 期　　　　　　　　　　　　　植　　　物　　　研　　　究 2003年　7 月
Vol.23　No.3　　　　　　　　　　　BULLETIN OF BOTANICAL RESEARCH July , 　2003
不同年龄兴安落叶松树干呼吸及其与环境因子关系的研究
姜丽芬1　石福臣2　祖元刚1　王文杰1　小池孝良3
(1.东北林业大学森林植物生态学教育部重点实验室 ,哈尔滨　150040)
(2.中国科学院地理科学与资源研究所 ,北京　100101)
(3.日本北海道大学演习林 ,札幌　060-0809)
摘　要　采用动态红外气体分析法研究了两个不同年龄兴安落叶松(Larix gmel inii Rupr.)人工
林内落叶松树干呼吸速率的季节变化 ,并分析了树干呼吸速率与环境因子的关系。两个年龄落叶
松树干呼吸速率均是从春季到夏季逐渐升高 ,高峰值出现在 7月(成熟林)和 8月份(幼林),之后
明显下降 。幼林落叶松的树干呼吸速率(变化范围是 1.99 ～ 6.15 μmol·m-2·s-1)显著高于成熟
林(变化范围是 1.52 ～ 3.38μmol·m-2·s-1)(P<0.05)。树干温度对树干呼吸影响较大 ,树干呼
吸速率与树干温度呈指数相关关系;成熟林和幼林树干呼吸的 Q10值分别为 1.96 和 3.44。当空
气相对湿度较低时 ,树干呼吸速率与其关系无明显规律 ,但当空气相对湿度很高时 ,能大大促进树
干的呼吸作用 。
关键词　兴安落叶松;树干呼吸;树干温度;Q10;空气相对湿度
STUDYON STEM RESPIRATIONOF LARIX GMELINII
OF DIFFERENT AGES AND ITS RELATIONSHIP
TO ENVIRONMENTAL FACTORS
JIANG Li-Fen 1　SHI Fu-Chen 2　ZU Yuan-Gang1
WANG Wen-Jie 1 KOIKE Takayoshi3
(1.Key Laboratory of Fo rest Plant Ecology , Ministry of Education , Northeast Forestry University , Harbin　150040)
(2.Institute of Geographical Sciences and Natural Resources Research , Chinese Academy of Science , Beijing　100101)
(3.Hokkaido University Forests , Sapporo　060-0809　Japan)
Abstract　By using a dynamic IRGA method , seasonal changes of stem respi ration rate in tw o larch
plantations w ith dif ferent ages w ere studied.Relationship between stem respi ration rate and environ-
mental factors were analy zed.Stem respiration rates of larches of bo th ages increased during spring
and summer , and peaked in July (the mature forest)and August (the young fo rest).During au-
tumn , stem respiration rates declined sharply .Stem respiration rate of the young larch stand , ranged
from 1.99 to 6.15 μmol·m-2·s-1 , was significantly higher than that of the mature larch stand ,
ranged from 1.52 to 3.38 μmol·m -2·s-1(P <0.05).There was an exponential correlation betw een
stem respiration rate and stem temperature.The temperature coefficient Q 10 values for stem respira-
tion of the mature fo rest and the young forest were 1.96 and 3.44 respectively.There w as no obvious
regularity betw een stem respirat ion rate and relative humidity at the lower atmospheric relative hu-
midity ,how ever , stem respiration w as apparently promoted at higher relative humidity range.
基金项目:本研究由中国科学院地理科学与资源研究所创新工程研究项目(CXIOG-E01-06)和(SJ10G-D01-02)及国家自然科学基金项目(30070133)资助。第一作者简介:姜丽芬(1971—),女 ,讲师 ,主要从事植物生理生态学研究 。收稿日期:2002-01-23
Key words　Larix gmelinii;stem respi ration;stem temperature;Q10;atmospheric relative humidi-
ty
　　Since 1950s , the concentration of atmospheric
g reenhouse gases has been sharply increasing because
of human activi ties and the g lobal climate change has
already become the most primary and direct change
of global change [ 1] .The global change of ecological
environment including the increase of CO2 concen-
tration and aggravating of global w arming has g reat-
ly threatened life system on the earth including hu-
man being [ 2] .“ The g reenhouse effect” and global
climate change have become the focus for people to
pay attention to.Accordingly it is one of the most
popular fields in modern ecology .
Forest ecosy stem has a very impo rtant signifi-
cance to carbon balance and global climate change
because 80% of aboveg round organic carbon on the
g lobal land and 40% of underground organic carbon
w as stored in forests[ 3 , 4] .According to Fang
Jingyun and Chen Anping , till 1998 , total forest
area in China w as 10 582×104 hm2and to tal carbon
storage in the forests was 4.75 Pg C.They suggest-
ed that in the recent over 20 years Chinese fo rests
played a role as sink of CO2w ith a mean annual se-
quest ration of 0.022 Pg C.This function of carbon
sink is mainly derived from the contribution of in-
creased refo restation and afforestation[ 5] .If the pro-
tection for the current forests is w ell done and refor-
estation is further reinforced , and together w ith the
maturity of middle and young aged forests w hich ac-
counted for very large propo rtion of total forest area ,
Chinese forests will be a huge potential carbon sink
in the future[ 6] .
Larch is widely distributed on eastern Eurasian
Continent and because of it s high capacity to f ix car-
bon and its longevi ty , larch fo rest is expected to be a
huge carbon sink[ 7～ 9] .In Northeast China , larch is
the most dominant species
[ 10] .However , info rma-
tion on the function of larch forests in Northeast
China as a carbon sink is very limited [ 9 , 11] .
With the dest ruction of natural forests , area of
man -made forests has been increasing .And in
1998 , the area and carbon of planted fo rests were
2311 ×104 hm2 and 0.72 Pg C respectively[ 5] .
Therefo re , we should know the CO2 sequest ration
capaci ty of these forests as a CO2 sink.However ,
there are no repo rts available about Larix gmelinii
plantations in Northeast China as a carbon sink [ 9] .
Stem respi ration in t rees results in substantial
CO2 release to the atmosphere.Stem respiration is
important because the bulk of the biomass in a ma-
ture fo rest stand is in woody tissue , and respi ration
in the living cells of this tissue occurs continuously ,
even trees are dormant [ 12] .Respiration of stem is
an important consti tuent of total respi ration in an e-
cosy stem.Woody tissues accounted for about 25%
of the total above-ground autot rophic respiration in
some forests such as in northern coniferous sites and
up to 50% of the above-g round autot rophic respi-
ration in some temperate deciduous forests [ 13 , 14] .
Stem respi ration could account for about 44% of a
community respiration , 58% of annual stand respi-
ration and 10% of an ecosystem respirat ion[ 15 ～ 17] .
Stem respiration would account for about 5% of
gross primary production (GPP)in a sw eetgum fo r-
est in eastern Tennessee , USA [ 12] and 10% of GPP
in tropical rain forests [ 18] .
It is likely that climate change can have great
ef fect on stem respiration because stem respi ration
has st rong response to temperature and CO2 enrich-
ment could increase 33% stem respiration[ 12 , 17 ～ 19] .
Although estimates of respi ratory loss for above
ground w oody t issue have been mainly conducted by
excision methods , these methods have been largely
superseded by various kinds of in situ measurements
[ 20] .Among these in situ measurements IRGA (In-
frared Gas Analyzer ) method prevails
[ 12 , 17 ～ 19 ,21～ 23] .
At present , research wo rk on stem respi ration
in our country is very few and for the very few re-
search work about stem respiration available , the
method applied w as static alkali absorption
method
[ 16] .Because there is a measure error that is
caused by non -complete absorption of CO2 , the
2973 期　　　　　　　　　　姜丽芬等:不同年龄兴安落叶松树干呼吸及其与环境因子关系的研究
static method to measure soil respiration has no t
been adoptedin foreign countries[ 24] .
In present research w ork , we use IRGA
method to measure the respiration rate of stem in or-
der to examine seasonal change of stem respiration
rate of larches of dif ferent ages and also to examine
the relat ionship betw een stem respiration rate and
environmental factors.
1 Study site
The research w ork w as done at Lao Shan Ex-
perimental S tation in the Mao er Shan Forest Farm
belonging to Northeast Forest ry University.This
station locates in the northwest part of the Zhang-
guangcai Mountains that belong to the Changbai
M ountains chain (45°20′N and 127°34′E , ca.340
m a.s.l.).Lao Shan Experimental Station belongs
to continental monsoon climatic zone.There are
typical four seasons usually w ith long w inter and
short summer.Winter is dry and cold w hile summer
is humid and hot.Precipitation is concentrated main-
ly in July and August.There is strong evapo ration
and very lit tle precipitation in spring w ith strong
wind resulting in dry climate.In autumn , the
weather condition is stable.Soil is typical dark -
brow n fo rest soil.Mean annual ai r temperature is
about 2.8℃ and mean annual ai r humidity is ca.
70%.Mean annual precipi tation and mean annual e-
vaporation is 723.8 mm and 1 093.9 mm respective-
ly.Mean annual sunshine time is 2471.3 hours.
Frost-free time is about 120 ～ 140 day s.
2　Materials and methods
Two stands of larch plantations w ere selected.
One is 31 years old (mature forest thereaf ter)and
the o ther is 17 years old (young forest thereaf ter).
These tw o stands w ere in the same sites.The basic
information about these tw o stands w as showed in
Table 1.
Table 1 The dif ferences between two stands of larch plantations
S tand Age Mean diameter(cm) Mean tree heigh t(m)
Density
(Individuals·hm-2)
Matu re forest 31 16.4±4.8 14.2±3.6 1420
Young forest 17 10.2±4.6 10.1±2.3 1533
We used the similar method described by Xu et
al.[ 17] and Wang et al.[ 23] to measure stem respira-
tion.This method ex tends the function of the Li-Cor
LI-6400-09 soil chamber to measure stem respira-
tion.The chamber w as fastened to the stems by a
cord or a tripod while measuring and the rubber plas-
ter w as used to seal the space between the chamber
and the stem.The chamber w as connected to a LI-
6400 po rtable photosynthesis sy stem(Li-Cor , Inc.,
Lincoln , NE)to analy ze gas exchange rate.The Li-
cor 6400 portable pho tosynthesis system can measure
several envi ronmental factors at the same time in-
cluding atmospheric relat ive humidity.Stem temper-
ature w as measured simultaneously by thermometers
RT-21S(ESPEC MIC CO RP ., Japan).
The stem respiration rates of larches were mea-
sured monthly , i.e., in the middle of every month
during g rowing seasons , we chose a typically fine day
to measure.In each stand we measured stem respira-
t ion rate of 5 larches and for each tree w e measured
three cycles.These 5 trees w ere chosen randomly ,
but thei r diameters w ere all about the mean diameter
level of the w hole stand.We measured respiration on
stems at a height of about 1.3 m above the g round.
The same trees were sampled throughout the study.
In order to minimize erro r caused by different
time of measurement , stem respiration rate of larches
of tw o stands w as measured almost in the same days
and the time interval of measurement between the
two stands was reduced as lit tle as possible.
3 Results
3.1　Seasonal changes of stem respi ration rate of
larches
298 　　　　　　植　　物　　研　　究　　　　　　　　　　　　　　　　　　23 卷
Fig.1　Seasonal changes of stem respiration rate of larches
in two larch plantations
　　The stem respi ration rate of larches in two plan-
tations in different months was show ed in Figure 1.
Stem respiration rate of larches of both ages in-
creased during spring and summer , and peaked in Ju-
ly (the mature forest)or August(the young fo rest).
During autumn , stem respiration rate declined
sharply.Stem respiration rate of the mature forest
ranged from 1.52 to 3.38μmol·m-2·s-1while stem
respiration rate of the young fo rest ranged from 1.99
to 6.15 μmol·m-2·s-1.
Fig.2　Stem temperature of the two larch
plantations in different months
Throughout the whole g rowing season , stem
respiration rate of larches of the young forest w as sig-
nif icantly higher than that of the mature forest(P <
0.05).The biggest dif ference appeared in Septem-
ber.In September , the stem respiration rate of
larches of the young forest w as more than tw ice
higher than that of the mature forest.The smallest
difference appeared in May.
3.2　Relationship between stem respiration and en-
F ig.3　Temperature response curves of stem
respiration of two stands
★Mature forest　 Young forest
vironmental factors
3.2.1 　Relationship between stem respi ration and
stem temperature
The stem temperature w as show ed in Figure 2.
The maximal value of stem temperature occurred in
July .The response of stem respiration of two differ-
ent fo rests to the stem temperature w as similar.In
general , with the increase of the stem temperature ,
respiration rate of the stem rised.However , there
w as an exception , namely , in August , although the
stem temperature w as much lower than that in July ,
stem respiration rate of young forest reached the peak
and stem respiration rate of the mature forest was al-
most the same w ith the peak value in July.This
probably w as caused by the higher atmospheric rela-
t ive humidity in August.
Fig.4　Seasonal variation of stem respiration
and relative humidity of two stands
To test the relationship between stem respira-
t ion rate and stem temperature , we applied an expo-
2993 期　　　　　　　　　　姜丽芬等:不同年龄兴安落叶松树干呼吸及其与环境因子关系的研究
nential function of the form y =β0eβ1 T , where y is
the measured stem respirat ion rate , β0and β1are fit ted
constants , and T is measured stem temperature.
The Q10 values , the indicator of temperature sensi-
tivi ty , were calculated as:Q10= e10β1.Exponential
curves of reg ression analyses are show ed in Figure 3.
Among all environmental factors , the stem tempera-
ture w as the most important factors to affect the
stem respi ration because exponential functions relat-
ing stem respirat ion to stem temperature accounted
for 75.6% of the seasonal variation in stem respira-
tion of the mature forest and 79.1% of the seasonal
v ariation in stem respiration of the young fo rest.The
temperature coef ficient Q 10 values for stem respira-
tion of the mature forest and the young forest w ere
1.96 and 3.44 respect ively .
3.2.2 　Relationship between stem respiration and
atmospheric relative humidity in the forests
F igure 4 show ed that there w as no st rong corre-
lation between stem respirat ion rate and atmospheric
relative humidi ty (correlation coefficient R2 = 0.44
for the mature forest and R2= 0.24 for the young
forest).But when the relative humidity of the air
w as very high , respiration of the stem could be
g reatly promoted.Fo r example , in August , the stem
temperature w as not the highest , but stem respira-
tion rate of the young forest reached the peak and
stem respiration rate of the mature fo rest w as almost
the same wi th the maximal value in July .This phe-
nomenon probably resulted f rom the highest relative
humidity of the ai r in August.
4 Discussion
The stem respi ration rate of the young larches
w as obviously higher than that of the mature larches.
This is probably because the life activi ty of organs of
the young trees is mo re vigorous than that of the ma-
ture t rees.Young trees are at the stage of vigorous
g row th and this needs mo re energ y.I t is stronger
respiration that can offer more energy fo r the vig or-
ous grow th.In the previous research wo rk , S tock-
fors and Linder had concluded that stem respiration
w as positively co rrelated with g row th rate
[ 25] , and
our results ag ree with that.
Xu et al.had pointed out that the diurnal and
seasonal variations in respi ration rate correlated well
w ith the corresponding stem temperature variation
[ 21] and our result w as consistent wi th it.Our result
w as also very consistent w ith a research result in
Japan [ 23] .
Although the response of stem respi ration of two
dif ferent fo rests to the stem temperature w as similar ,
there w as a difference could be noted by using corre-
lation analy sis.Both the values of co rrelation coeffi-
cient betw een stem respi ration and stem temperature
and Q 10value of young fo rest w ere higher than those
of the mature forest.This indicated that the response
of stem respiration of young trees to stem tempera-
ture w as more sensitive than that of mature t rees.
The possible reason for this phenomenon might be
that the activi ty of org ans of the mature trees w as
relatively stable while the activity of org ans of the
young trees w as more vigorous.
The temperature response of w oody tissue respi-
ration tends to have a Q10 betw een 1.5 and 2.5 [ 26] .
Q10 value of the mature forest (1.96)was good
w ithin this range , but Q 10 of the young forest (3.
44)was much higher than the maximal value of this
range.While Q 10of the mature fo rest was similar to
Q10 of sw eetgum trees in eastern Tennessee , USA
(Q 10 =1.94 and 2.12)[ 12] , lower than Q 10 of bal-
sam fi r stands in western Newfoundland and New
Brunsw ick(Q 10=1.96 ～ 2.48)[ 19] , and low er than
Q10 of young ponderosa pines in the Sierra Nevada
M ountains , California (Q10 varied f rom 2.4 to 2.9
among the t rees)[ 17] , Q 10 of the young forest w as
higher than all of them.Bo th Q10values of the ma-
ture forest and the young forest were higher than
Q10 of tw o t ropical rain forests (Q10= 1.8 and 1.
6)[ 18] .
As for the response of stem respiration to the
relative humidity of the ai r , there were no relevant
reports available.In this study , when relative hu-
midity of the air w as low , i t didn t have obvious ef-
fect on stem respi ration , but in August , because of
f requent rainfall , the relative humidity of the air w as
very high and the stem respiration of larches of both
300 　　　　　　植　　物　　研　　究　　　　　　　　　　　　　　　　　　23 卷
ages w as g reatly enhanced.In other wo rds , high hu-
midity of the air af fected stem respi ration w hile low
humidity didn t.One of the possible reasons w hy
the high relative humidity of the air promo tes respi-
ration of the stem may be that it accelerates the de-
composition of the dead bark.The further study w as
needed to explain the reason why high relat ive hu-
midity could promote stem respiration of larches.
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3013 期　　　　　　　　　　姜丽芬等:不同年龄兴安落叶松树干呼吸及其与环境因子关系的研究