全 文 ：松嫩草原拂子茅种群热值 、生物量和能量动态的研究
郭继勋1　王若丹2　王　娓1
(1.东北师范大学草地研究所 ,长春 130024;
2.中国科学院生态环境研究中心 ,北京 100093)
摘要:　拂子茅(Calamagrostis epigejos)全株和叶的热值的季节变化规律相似 ,呈双峰曲线 , 2 个峰值分别出现在 6 月
中旬和8 月初。茎热值的变化呈单峰曲线 ,峰值出现在8 月初。穗和立枯体的热值变化呈波动型 , 穗的最大值出现
在9 月初 ,立枯体出现在 6 月中旬。地上部能量现存量的季节变化与生物量变化同步 , 呈单峰曲线。不同季节能量
在各器官中分配比率为:6月中旬为叶>茎>穗;7月初为叶>茎>立枯体>穗;7 月中旬为叶>茎>穗>立枯体;8
月至 9月为叶>茎>立枯体>穗。能量的垂直分配 ,地上部为从地表至 20 cm 高度逐渐增加 , 最大值在 10 ～ 20 cm
层占地上部总能量的 26.91%,然后逐渐下降。地下部能量垂直分布规律是随着深度增加而逐渐减少 , 能量集中分
布在地下 0～ 10 cm 层 , 占地下部总能量的 69.01%。
关键词:　拂子茅;热值;能量;生物量
中图分类号:Q947.5　　　文献标识码:A　　　文章编号:0577-7496(2001)08-0852-05
Study on Dynamics of Calorific Value , Biomass and Energy in Calamagrostis
epigejos Population in Songnen Grassland
GUO Ji_Xun1 , WANG Ruo_Dan2 , WANG Wei1
(1.Institute of Grassland science , Northeast Normal University , Changchun 130024 , China;
2.Research Center for Eco_environment , The Chinese Academy of Sciences , Beijing 100095 , China)
Abstract:　There were two peaks of seasonal changes of the calorific value in shoot and leaves of Calama-
grostis epigejos in middle June and in the early August respectively.The calorific value in stem presented a sin-
gle peak curve which appeared in the early August.The calorific values in inflorescence and dead standing
showed a fluctuation and the peak value of inflorescence was in the early September and that of dead standing
was in middle June.The seasonal changes of energy standing crop on the above_ground part synchronized with
that of the biomass , which presented a single peak curve.The energy allocated to each organ in different sea-
sons was in the order as leaves >stem >inflorescence in middle June , leaves >stem >dead standing >in-
florescence in early July , leaves >stem >inflorescence >dead standing in middle July , and leaves >stem
>dead standing >inflorescence from August to September.The vertical allocation of energy in the parts of
above_ground was that the energy value gradually increased from the surface to the 20 cm high level and the
maximum value at the 10-20 cm high level which made up 26.91%of energy on the above_ground partion ,
and then it was decreased.In the under_ground portion , the energy value progressively decreased with depth
and the maximum value was at 0-10 cm depth layer which made up 69.01% of energy of the under_ground
portion.
Key words:　Calamagrostis epigejos;calorific value;energy;biomass
　　 The research on the dynamics of calorific value ,
biomass and energy of grass as well as their relationship is
the basis for assessing energy immobilization , conversion
and utilization efficiency and nutrient value of the grass.
Energy is a driving force for normal operation of the
ecosystem.Therefore , it is of great theoretical signifi-
cance to discover the formation mechanism and ecological
process of the biomass by investigating the changing rule
of calorific value , energy and spatial distribution of the
plant.So far , relevant researches in the grassland ecosys-
tem at home and abroad mainly include:dynamics of en-
ergy accumulation of the main communities in different
types of the grasslands , the increasing ratio of standing
energy crop and the energy conversion ratio[ 1-3] ;
Received:2000-10-30　Accepted:2001-02-28
Supported by the National Natural Science Foundation of China(39970537).
植　物　学　报　2001 , 43(8):852-856 　 　 　 　 　 　 　
Acta Botanica Sinica
dynamics of the biomass and energy allocation[ 4-5] ;grass-
land productivity structure , energy flow and energy dy-
namic model[ 6-7] ;calorific value , energy determination
and dynamic analysis of the main grassland plants[ 8-10] ;
the relationship between calorific value and chemical com-
position and nutrient substance of plants , the effect of
grazing on calorific value , biomass and energy of the
plants[ 11-12] .Calamagrostis epigejos is the main grass
grow abundantly with extensive distribution in Songnen
Grassland.It has played an important role in regulating
the microclimate and keeping the stability of the structure
and function of grassland ecosystem.C.epigejos has a
high productivity and good quality which provides a high
nutrient value for animals feeding.This paper discussed
the dynamics of calorific value , biomass and energy as
well as their relationship.The research also provided a
theoretical basis for rational utilization of grassland re-
sources and improvement of the conversion of the material
and the energy between grass and livestock.
1　Study Area and Methods
1.1　Study area
The study was conducted in Changling County(123°
44′-47′E , 44°40′-45′N)Jilin Province , China.It
enjoys a typical continental climate with annual rainfall
amounting 470.6 mm , which mainly distributed in June
-September.The annual evaporation amount 1 600.2
mm.The annual average temperature is 4.9 ℃with the
warmest weather averaged 23.2 ℃in July and the coldest
weather in January (-16.4 ℃ in average).The area
belongs to meadow grassland in which Calamagrostis
epigejos is extensively distributed.There are a high satu-
ration value of 14 m
-2
and a diversity index of 1.628 in
the C.epigejos community.In addition , the degree of
coverage , density and height of C.epigejos dominate in
the community with a biomass of 60%-80% of the total
biomass of the community.
1.2　Methods
The study was carried out in 1996 , at which time the
annual air temperature was 5.8 ℃, rainfall was
307.0 mm , evaporation capacity was 1 611.1 mm and
daylight hour was 2 624.8.The plot area was selected in
a typical C.epigejos community.The experimental area
included 3 replicate subplots.C.epigejos turns green
from late April to early May , goes into heading in middle
June , reaches the climax of biomass in the middle August
and ceases growing in September.In order to determine
the calorific value of different organs and biomass and the
allocation of energy in different organs , the above_ground
parts of the plant was sampled from June to September in
half a month interval.The experimental area included 5
replicate 1 m2 subplots.The above_ground parts of C.
epigejos in the community were cut down entirely and oth-
er kinds of plants were all removed.The collected sample
was cut every 10 cm to make every section from the basal
portion to the top.The sections were divided into stem ,
leave , inflorescence and dead standing.The sampling
time of the below_ground parts was arranged at the begin-
ning of August.The below_ground parts were cut every 10
cm depth until 50 cm.The area of subplot was 1/4 m2.
Mud and messes were washed off the plant.Then the
samples were dried at 80 ℃until constant weight to de-
termine the biomass.The calorific value was determined
by TA-2000 thermal flowmeter.Finally the energy value
was calculated.
2　Results and Analysis
2.1　The seasonal dynamics of calorific value of C.
epigejos
The seasonal dynamics of calorific value of plants re-
late to the substances containing the energy and biological
character of the plants.C.epigejos is a relatively high
plant and its stem functions as a supporting organ of the
plants.Therefore , the calorific value of the stem steadily
increased from the initial period of growth to the climax
period (August).During the period , the nutrient was ac-
cumulated to ensure the growth of the stem.The seasonal
dynamics of calorific value showed a single_peak curve
and the peak value appeared at the beginning of August.
Leave functions as a photosynthetic organ.At the
initial period of growth , a large amount of high_energy
substances was accumulated to support the leaf growth.
During the period , the calorific value was high.From
middle June to middle July , which was the heading stage
of C.epigejos , the plants entered the stage of reproduc-
tion , when the nutrient transferred to the reproductive or-
gan.During the period , the calorific value was in a de-
clining trend.Up to August , the plant ceased its repro-
ductive growth and the calorific value began to increase.
In the final stage of the growth , the nutrient transferred to
the below_ground parts of the plants resulting in a declin-
ing calorific value.The seasonal dynamics of calorific val-
ue showed a two_peak curve with the peak values in the
middle June and at the beginning of August.
From middle June to the beginning of July , a large
amount of the energy_containing substances were stored to
form flowers.The calorific value was high.From middle
July to the beginning of August , the plants blossomed ,
8 期 郭继勋等:松嫩草原拂子茅种群热值 、生物量和能量动态的研究(英) 853　
which consumed the energy and prepared for the develop-
ment of seeds , so the calorific value was low.With the
accumulation of the substances containing the energy in
the seed , the calorific value of inflorescence increased
slowly .In comparison with other organs , the calorific val-
ue of inflorescence was the highest.During the growth pe-
riod , the calorific value slightly fluctuated.The maximum
value appeared at the beginning of September.
Dead standing appeared at the beginning of July.
The calorific value was low during the growth period.A
seasonal alteration calorific value was seen , with a maxi-
mum value at the beginning of July.During the growth
period , the changing rule of average calorific value in dif-
ferent organs was inflorescence >leave >stem >dead
standing.
The seasonal dynamics of calorific value of the plant
are organ dependent.In C.epigejos , the calorific value
was highest in the initial growth period , declined gradual-
ly.Then a second peak appeared in the climax period of
growth(at the beginning of August)that declined gradu-
ally afterwards , being lowest in September(Table 1).
Fig.1.　The seasonal changes of the calorific value , biomass and
energy of Calamagrostis epigejos population.
2.2　The seasonal change of energy and its allocation
in different organs and biomass in the above_ground
parts of C.epigejons population
The seasonal dynamics of energy standing crop in
C.epigejons population depends on the change of
biomass and calorific value.The energy standing crop e-
quals the biomass multiplied by calorific value.The sea_
sonal change of the above_ground biomass of C.epigejons
population showed a single_peak curve with the maximum
in the middle August , 443.33 g·m2.In the final stage of
growth , the calorific value declined gradually.The sea-
sonal change showed a steady trend with little variation.
The seasonal dynamics of energy standing crop was consis-
tent with the changing rule of biomass.The energy was
gradually accumulated to a peak value , 8 282.20 KJ·m2
from the initial period of growth to middle August.Then
the energy standing crop declined(Fig.1).The seasonal
change of energy standing crop depended mainly on the
above_ground biomass because the accumulation process of
biomass synchronized with that of energy.The change of
calorific value had little effect on that of the energy be-
cause the differences of calorific value of the whole plant
were far lower than that of the above_ground biomass.
In different growing seasons , the allocation ratio of
energy in the same organ was different.The allocation ra-
tio of energy in the stem showed an increasing trend from
the initial growth period to the middle of July with the
maximum reaching 41.74% of the total energy standing
crop of the above_ground.The allocation ratio kept a
steady trend from middle July to middle August.The allo-
cation ratio was more than 40%.Up to September , it de-
clined rapidly.The allocation ratio of energy of the leave
basically showed a declining trend with the maximum in
middle June , 50.41%of the energy standing crop of the
above_ground.The allocation ratio varied between 48%
and 41%from the beginning of July to the middle of Au-
gust.The allocation ratio of the inflorescence showed a
fluctuating tendency with the maximum in middle June ,
14.5%of the above_ground standing crop of the energy.
The allocation ratio varied between 6.4% and 7.7%
from early July to middle August.In September , the allo-
cation ratio declined sharply to 2.92%.The allocation
ratio of dead standing showed basically an increasing
trend.It changed slightly between 6% and 9% from the
beginning of July to the middle of August.After middle
August , the allocation ratio of dead standing increased
rapidly due to the high death rate of the plants.At the
beginning of September , the allocation ratio reached
28.97%(Fig.2).
Table 1　The seasonal changes in calorific values of Calamagrostis epigejos(M±SE , n=15)
Time Stem Leaves Inflorescence Standing dead Shoot
6/ 15
7/1
7/ 15
8/1
8/ 15
9/1
1 9554.04±3.56
1 9733.78±6.32
1 9750.50±5.46
2 0186.47±9.32
1 8404.54±5.39
1 7505.84±4.25
2 1873.94±6.53
1 9386.84±3.45
1 8024.16±4.96
2 0372.75±5.47
1 9370.12±7.24
1 7869.54±8.03
2 1675.26±3.47
2 1581.34±7.98
2 0770.42±6.31
2 0848.03±5.78
2 1209.32±3.99
2 1873.94±4.56
1 8312.84±2.79
1 6101.36±4.36
1 8136.29±5.12
1 5812.94±6.45
1 6887.20±2.91
2 0972.83±2.89
1 9891.20±4.32
1 8731.18±7.56
2 0099.86±9.12
1 8681.80±5.55
1 6827.40±5.23
854　 植　物　学　报　　Acta Botanica Sinica 43 卷
Fig.2.　The allocation ratio of energy in different organs.
　　There existed some seasonal variations of allocation
ratio of the energy among different organs.The changing
rule appeared to be in the order in the middle of June ,
leave >stem >inflorescence;at the beginning of July
leave>stem>dead standing>inflorescence;in the mid-
dle of July , leave>stem>inflorescence>dead standing;
during August and September , leave >stem >standing
dead>inflorescence.
2.3　Vertical allocation of energy of C.epigejons
population
The energy allocation in the part of above_ground
and below_ground of C.epigejos population in the early
August , a heavy growth season , was analyzed.During
that period , the total energy standing crop of C.epigejos
population was 33 208.57 KJ·m2.The energy in the part
of the above_ground was 7 024.90 KJ·m2 , 21.15% of
the total energy standing crop and the energy in the part of
the below_ground was 26 183 KJ·m2 , 78.85%of the to-
tal energy standing crop , which was 3.73 times than that
of the above_ground.
The energy structure tower in the part of the above_
ground was divided into 11 layers.The energy located in
layer 0-10 cm (from soil surface to 10 cm high)was
24.66% of the total energy in the part of the above_
ground.The energy located in layer of 10-20 cm was
highest , 1 890.10 KJ·m2 , 26.91% of the total energy in
the part of the above_ground.The energy decreased with
the degree of the height within 20-70 cm.The energy
located in the layer of 20-30 cm , layer of 30-40 cm ,
layer of 40 -50 cm and layer of 50 -60 cm was
22.42%, 10.09%, 6.72%, 2.24% and 1.35% of
the total energy in the part of above_ground respectively.
The layer of 0-30 cm was made up of stem , leaves and
dead standing , that of 30-50 cm was made up of stem
and leaves , 50-70 cm was made up of stem , and the
layer of 70-110 cm was made up of inflorescence only ,
which was the characteristic of the energy structure tower
of C.epigejos population.The energy located in the in-
florescence in the layer of 70-110 cm was 5.61%of the
energy in the part of the above_ground(Fig.3).
Fig.3.　The vertical allocation of energy of Calamagrostis epigejos
population.
The vertical structure in the part of the below_ground
of C.epigejos population was divided into 5 layers.A
layer was determined every 10 cm depth from soil surface.
The vertical allocation of energy standing crop de-
creased gradually with the degree of the depth.The ener-
gy distributed intensively in the layer of 0-10 cm with
the standing energy croup 18 068.30 KJ·m2 , 69.01%of
the total energy standing crop in the part of the below_
ground because of the densely distributed root system in
this layer.The energy value located in the layer of 10-
20 cm , 20-30 cm , 30-40 cm and 40-50 cm was
16.08%, 7.57%, 4.73% and 2.61% of the total en-
ergy in the part of the below_ground respectively.
3　Discussion
The seasonal changes of calorific value and energy of
plants related to the biological character and biotempera-
ture rule of the plants.The change of calorific value in
different organs was essentially caused by different content
of the energy containing substances in different organs.
The seasonal change of energy standing crop depended on
the utilization ratio of photosynthesis of the plants and ac-
cumulation ratio of biomass.The energy allocation ratio
depended on the growth and development rule and repro-
ductive strategy.So far as the perennial grass conducting
vegetative reproduction was concerned , the energy value
allocated in the leaves and stem was higher than that in
the inflorescence.In the final growing period , the energy
was transferred to the below_ground and accumulated to
ensure the growth , development and reproduction of the
plants.Therefore the energy allocated in the part of the
below_ground was far more than that in the part of the
above_ground.
8 期 郭继勋等:松嫩草原拂子茅种群热值 、生物量和能量动态的研究(英) 855　
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856　 植　物　学　报　　Acta Botanica Sinica 43 卷