全 文 ：第 28 卷 第 1 期 植 物 研 究 2008 年 1 月
Vol． 28 No． 1 BULLETIN OF BOTANICAL ＲESEAＲCH Jan．， 2008
Foundation item:The National Natural Science Foundation of China (Grant No． 30471423) ，Primary Ｒesearch Project of Great Basal Ｒesearch of
Science and Technology Ministry (Grant No． 2004CCA03000) and the National Natural Science Foundation of Inner Mongolia (Grant No．
200408020506)
Author introduction:HONG Yu(1969—) ，female，Ph． D． The research field is protection biology．
Ｒeceived date:2007 － 05 － 24
臭柏群落在不同演替阶段某些群落特征的研究
红 雨1 王林和2
(1．内蒙古师范大学生命科学与技术学院，呼和浩特 010022)
(2．内蒙古农业大学，呼和浩特 010022)
摘 要 毛乌素沙地臭柏群落自发演替的基本过程为:半流动沙地沙竹 +沙米(有时是白沙蒿)
群落→半固定沙地油蒿 +臭柏群落→固定沙地臭柏 +硬质早熟禾群落→固定沙地臭柏群落→老
固定沙地臭柏 +苔藓群落，最后可能发展成为地带性的本氏针茅草原。演替早期向演替亚顶级群
落发展过程中物种多样性逐步增加，生态环境逐渐变湿。随着臭柏的衰退，某些喜湿植物的退出，
物种多样性开始减少，群落生境向旱生化发展。
关键词 群落特征;臭柏;毛乌素沙地;演替
中图分类号:Q948 文献标志码:A 文章编号:1673 － 5102(2008)01 － 0109 － 05
Community Characteristics of Sabina vulgaris Community
under Different Stages of Succession
HONG Yu1 WANG Lin-He2
(1． College of Life Science and Technology，Inner Mongolia Normal University，Huhhot 010022)
(2． Inner Mongolia Agricultural University，Huhhot 010022)
Abstract The autogenic succession process of vegetation in Mu Us Sandland is from Psammochlou uil-
losa + Agriophyllum arenarium (Artemisia sphaerocephala)on semi-drift-sand dunes to Artemisia ordosica
+ Sabina vulgaris on semi-fixed-sand dunes，then through S． vulgaris + Poa sphondylodes on fixed-sand
dunes and S． vulgaris on fixed dunes of subclimax stage to S． vulgaris + moss community on old-fixed-
sand dunes，which maybe pass to a Stipa bungeana steppe finally． The S． vulgaris community succession
contains progressive and regressive succession． Species diversity increased with the development of the
progressive succession，and the ecological environment began getting wet． Along with the declining of S．
vulgaris，some plants exited，species diversity decreased and the habitat of community began getting dry．
Key words community characteristics;Sabina vulgaris;Mu Us Sandland;succession
Mu Us Sandland，with an altitude of about 1 300
to 1 600 m，is situated in the south of Erdos altiplano．
Its mean annual temperature is about 6 to 9℃ ． July is
the hottest month in which the mean temperature is a-
bout 20 to 24℃，In January，the mean temperature is
about － 12 to － 8℃ which is the coldest month． The
mean annual precipitation ranged from 250 to 490 mm
which descending from southeast to northwest． Mu Us
Sandland is a ecotone which has special geographical
landscape［1］． Sabina vulgaris，belonging to Cupres-
saceae，Cypress Family，is a naturally unique ever-
green coniferous herb in Mu Us Sandland． Its root
system is strong，and branches and leaves are flourish
which can also bear wind and sand． It is a excellent
and rare tree which can fix sand and protect soil． Un-
der the diagusting sandy environment，the coverage of
S． vulgaris can reach 70% to 95%，forming a single
dominant community［2］． Many scholars at home and
abroad have paid more attention to S． vulgaris be-
cause of its advantages． This study discussed the sta-
ges of succession of natural S． vulgaris community
and its adaptability to the sandy environment under
different stages of succession in Mu Us Sandland，
which may provide basic theory and technique for the
resuming of the damaged ecosystem．
1 Materials and methods
This study was carried out in the north experi-
mental area of Mu Us Sandland．，The stages of suc-
cession were chosen through space instead of tempo-
ral［3 ～ 5］． First，we selected a belt transect in the
field． Length and width of belt transect are 2 000 m
and 4 m，respectively，and sample plots with an area
of 4 m × 4 m is sequentially obtained in the bel tran-
sect． Second，we investigated the sample plots and
the acquired data of community，population and soil
was processed using cluster analysis，quantity analysis
was conducted by PCA，DCA，TWINSPAN． Combi-
ning the above results and succession rule，the stages
of succession were divided． Finally，15 sample plots
(4 m ×4 m)in each stage of succession were random-
ly chosen in north experimental area in June and Au-
gust 2004，the community characteristics under differ-
ent stages of succession were compared and analyzed．
2 Ｒesults and analysis
2． 1 Succession of S． vulgaris community
In Mu Us Sandland，the soil surface has no veg-
etation and the sunlight is strong，the wind speed is
fast and the sand tends to dry and drift easily，many
plants can not survive，we could only find some pio-
neer plants such as Agriophyllum arenarium，Pugio-
nium cornutum，Corispermum patelliforme，Psammo-
chloa uillosa and Artemisia sphaerocephala． Because
of the occupancy of the pioneer plants，the fluid of
sand dune was weakened，the sand was dominated by
these plants，forming P． uillosa + A． arenarium (A．
sphaerocephala)community on semi-drift-sand dunes．
In such environment，Artemisia ordosica and S． vul-
garis survived well，resulted in an increase of the
density and coverage of vegetation，more soil moisture
was absorbed． At the same time，the frequency and
degree of moisture stress of plant in community in-
creased，A． sphaerocephala began to die and A． ordo-
sica + S． vulgaris formed constructive species togeth-
er［6］，and then A． ordosica + S． vulgaris community
on semi-fixed-sand dunes formed． In Mu Us Sand-
land，the root system of S． vulgaris，which was stron-
ger than that of A． ordosica，was distributed like a
down-pyramid and mainly concentrated in the range of
0 － 60 cm soil layer where they use moisture and nu-
trition，and the fine roots of S． vulgaris were also
more than those of A． ordosica in the range of 30 － 50
cm soil layer［7］． When soil moisture could not satisfy
their requiring，A． ordosica declined and S． vulgaris
became the unique dominant species． Sand began to
be fixed，surface soil accumulated and organic matter
and nutrient of soil increased，which provided surviv-
al conditions to herbal plants． Cleistogenes spuarrosa，
Oxytropis glabra，Potentilla anserine，Clematis intri-
cate，Thalictrum squarrosum and Melilotus suaveolens
in arid grass began to come in and Poa sphondylodes
grew quickly，then S． vulgaris + P． sphondylodes
community on fixed-sand dunes formed． Because of
integrated effects by root system secretion of plants
and died organ，gravel was eroded and broken up，
died organ also made organic matter accumulated，
and then，sandiness soil grew up． At the same time，
soil moisture and fertility increased， which made
many hygrophytia， mesoxerophytic and mesophyte
plants came into the S． vulgaris communities (Table
1)． Generally，in this stage，the growth of S． vulgar-
is is the best and individuals are higher，breadth are
bigger，coverage and productivity are higher，then S．
vulgaris on fixed dunes of subclimax stage formed，
which is the most stable stage before Stipa bungeana
steppe which maybe climax community in this are-
a［1，6］． With the surface dust accumulated and crusts
became thick，surface organic matter broke up and
fertility increased，which were very important for the
011 植 物 研 究 28 卷
Table 1 The main types and basic characteristics of Sabina vulgaris community at different stages of succession
Latin name
Abundance
Ⅰ Ⅱ Ⅲ Ⅳ
Stage Ecological type
Sabina vulgaris cop1 ～ cop2 cop2 ～ cop3 cop2 ～ cop3 cop1 ～ cop2 Ⅰ，Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Ｒhamnus erythroxylon un un un un Ⅰ，Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Artemisia ordosica cop1 sp sol sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Clematis intricata sol sol Ⅱ，Ⅲ mesoxerophytic
Caragana korshinskii un un un Ⅰ，Ⅱ，Ⅳ xerophyte
Silene repens var． angustifolia sp sp sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Lespedeza davurica sp sp sol Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Oxytropis psammocharis sol sol sol sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Oxytropis glabra sol sol Ⅱ，Ⅲ mesophyte
Oxytropis giabra． var． tanus sol sol sol Ⅱ，Ⅲ，Ⅳ mesophyte
Potentilla anserina sp cop1 ～ sp sol Ⅱ，Ⅲ，Ⅳ mesophyte
Potentilla chinensis sol sp cop1 ～ sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Potentilla sericea sol sol sol sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Astragalus meliiotoides sol sol sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Polygala tenuifolia sol sp sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Euphorbia esula sol sol cop1 sp Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Bupleurum scorzonerifolium sol sol cop1 sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Limonium bicolor un un un 1，Ⅱ，Ⅲ xerophyte
Cynanchum komarovii sol un un sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Cynanchum thesioides sol sol sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Scutellaria baicalensis un sol sol Ⅰ，Ⅱ，Ⅲ mesoxerophytic
Plantago asiatica sol sp sp Ⅱ，Ⅲ，Ⅳ mesophyte
Plantago major sol sp Ⅱ，Ⅲ， mesophyte
Orobanche coerulescens sol sol sol Ⅱ，Ⅲ，Ⅳ xerophyte
Ｒubia lanceolata sol un sp ～ cop1 un Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Heteropappus alaicus sol sp sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Inula britanica sol sol Ⅱ，Ⅲ mesophyte
Artemisia argyi sol sol sol sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Sonchus arvensis sol sp ～ cop1 sol Ⅱ，Ⅲ，Ⅳ mesophyte
Carex duriuscula sol sol sp ～ cop1 Ⅰ，Ⅱ，Ⅲ，Ⅳ ophytic
Asparagus dauricus sol sol sol sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Pennisetum centrasiaticum sol sol sp Ⅰ，Ⅱ，Ⅲ mesoxerophytic
Psammochloa villosa sol Ⅰ xerophyte
Achnatherum splendens sol sol sp Ⅰ，Ⅱ，Ⅲ mesoxerophytic
Artemisia dalailamae cop1 sol sp Ⅰ，Ⅱ，Ⅳ xerophyte
Cleistogenes squarrosa sp cop1 sp sp Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Polygala sibirica sp sp sp Ⅱ，Ⅲ，Ⅳ xerophyte
Allium tenuissimum sol sol sol un Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Thalictrum squarrosum sp sol Ⅱ，Ⅲ mesoxerophytic
Glaux maritime sol Ⅲ mesophyte
Saussurea amara sp Ⅲ mesophyte
Allium polyrhizum sol Ⅲ xerophyte
Allium mongolicum sol sol cop1 sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Leymus chinensis cop1 Ⅲ mesoxerophytic
Taraxacum mongolicum sol Ⅲ mesophyte
Poa sphondylodes sol cop1 sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Halerpestes ruthenica un Ⅲ mesophyte
Vicia villosa sol sol sol Ⅰ，Ⅱ，Ⅲ mesoxerophytic
Ferula bungeana sol sol sol un Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Agropyron mongolicum cop1 sol sol Ⅰ，Ⅱ，Ⅲ xerophyte
Eqisetum hyemale sol sol cop1 Ⅰ，Ⅱ，Ⅲ mesophyte
Artemisia mongolica cop1 Ⅲ mesophyte
Artemisia sacrorum sol sol sp Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Phragmites australis sol Ⅲ hygrophytia
1111 期 红雨等:臭柏群落在不同演替阶段某些群落特征的研究
Contiune Table 1
Latin name
Abundance
Ⅰ Ⅱ Ⅲ Ⅳ
Stage Ecological type
Pulsatilla chinensis un Ⅲ mesophyte
Agriophyllum pungens sol Ⅰ mesoxerophytic
Plantago depressa sol cop1 ～ cop2 sp Ⅱ，Ⅲ，Ⅳ mesophyte
Melilotus suaveolens sp sol sp Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Puccinellia distans sol Ⅲ mesophyte
Elymus dahuricus un sp Ⅱ，Ⅲ mesophyte
Lappula myosotis sp sol sol sol Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
Medicago falcata sol sp Ⅱ，Ⅲ mesoxerophytic
Erodium stephanianum sol sol sol sol Ⅱ，Ⅲ Mesoxerophytic
Dracocephalum moldavi sp sp Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Euphorbia humifusa sp sol sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Echinops gmelini sol un Ⅰ，Ⅱ xerophyte
Leonurus sibiricus un sol sp Ⅰ，Ⅱ，Ⅲ mesoxerophytic
Artemisia annua sol cop1 sp un Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Artemisia scoparia sol cop1 sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesoxerophytic
Corispermum candelabrum sp sol sol Ⅰ，Ⅱ，Ⅳ psammophyte
Ixeris sonchifolia sol sol sp sol Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Bassia dasyphylla sp sol sol Ⅰ，Ⅱ，Ⅳ xerophyte
Eragrostis pilosa sol Ⅲ mesophyte
Chenopodium aristatum sol sol Ⅲ，Ⅳ mesophyte
Setaria viridis cop1 sp cop1 sp Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Chenopodium glaucum un sol Ⅱ，Ⅲ mesophyte
Chenopodium acuminatum sol un sol cop1 Ⅰ，Ⅱ，Ⅲ，Ⅳ mesophyte
Bidens parviflora sol Ⅲ mesophyte
Orostachys fimbriatus cop1 sp sp cop1 Ⅰ，Ⅱ，Ⅲ，Ⅳ xerophyte
growth of herbal plants of grass vegetation and moss
community． Those plants often appeared under the S．
vulgaris shrub then expanded gradually and could fix
crusts above the soil surface and prevented the precip-
itation，made the soil moisture of root system lacked
and influenced the growth of S． vulgaris． Moss com-
munity dominated and formed S． vulgaris + moss
community on old-fixed-sand dunes［3，4］． Table 1
showed the species numbers in forth stage decreased
(47 species)． Under this condition，S． vulgaris shrub
began to die，maybe pass to a S． bungeana steppe．
2． 2 Changes of community characteristics under
different stages of succession
2． 2． 1 Changes of community coverage，species
diversity and ecological types
The ascending and descending proportion of veg-
etation coverage were not the same and this asymmetry
reflected the changeability of each stage in succession
process． Less the ascending and descending propor-
tion，more this stage changeable，whereas，more in-
stable． From the ascending and descending proportion
of different stages，It was found that the third stage
was the most stable，but the forth one is the least．
Different stages of succession had influence on the
morphology of the plants． The plant height of S． vul-
garis decreased first，and then increased under differ-
ent stages of succession． The variance analysis
showed that the difference among stages was signifi-
cant． The change of growth mass was the same as the
plant height (Table 2)．
There are 80 different plants in four stages (Ta-
ble 1)． From Table 3，it can be found that species a-
bundance increased with the process of the succes-
sion． The species number was the most in the third
stage，reached 73，and then decreased in the forth
stage． From early stage to subclimax of succession，
the ecological environment became wet，the xerophyte
plants decreased and the mesophyte ones increased．
In the forth stage，some hygrophytia plants exited and
mesophyte plants decreased， whereas， xerophyte
plants increased，then the habitat of community be-
came dry (Table 3)．
211 植 物 研 究 28 卷
Table 2 Changes of community characteristics at different stages of succession
Ⅰ Ⅱ Ⅲ Ⅳ
Community coverage (%) 35 70 90 40
Coverage of Sabina vulgaris(%) 20 － 30 40 － 70 70 － 90 30 － 40
Growth mass (cm) 8． 47 ± 2． 92a 7． 21 ± 3． 44a 7． 91 ± 2． 89a 5． 17 ± 1． 71b
Plant height (cm) 55． 16 ± 16． 72b 58． 22 ± 9． 01ab 67． 26 ± 18． 45a 54． 68 ± 17． 97b
Numbers of species 48 65 73 47
Note:Different letters in the same colmn indicate significant difference (p ＜ 0． 05)
Table 3 Changes of ecological types of Sabina vulgaris community at different stages of succession
Succession stage Xerophyte Mesoxerophytic Mesophyte Hygrophytia
Ⅰ 23(47． 92%) 14(29． 17%) 11(22． 92%)
Ⅱ 22(33． 85%) 21(32． 30%) 22(33． 85%)
Ⅲ 19(26． 03%) 22(30． 14%) 31(42． 47%) 1(1． 36%)
Ⅳ 20(42． 55%) 12(25． 53%) 15(31． 92%)
2． 2． 2 Changes of β-diversity index
Generally，β-diversity index denotes comparabil-
ity index among communities or velocity of species at
different habitats in the same geography area［6］． It re-
flects the structure type， organic level， develop
stage，stability and difference of habitat between com-
munities and ecosystems［8，9］． Least the comparability
of species，lower the β-diversity index in different
habitats or different places in the same habitat．
Table 4 The β-diversity index at different stages of suc-
cession
Ⅱ Ⅲ Ⅳ
Ⅰ 0． 701 5
Ⅱ 0． 759 5
Ⅲ 0． 564 1
From Table 4，it can be found that changes a-
mong stages in the succession process are not the
same． The higher the comparability between the
neighbor stages was，the higher stability was，vice
versa． The order of stages of succession by stability
was S． vulgaris on fixed dunes ＞ A． ordosica + S．
vulgaris on semi-fixed-sand dunes ＞ S． vulgaris + P．
sphondylodes on fixed-sand dunes ＞ S． vulgaris +
moss community on old-fixed-sand dunes． During the
process of the vegetation succession in Mu Us Sand-
land，the third stage (S． vulgaris on fixed dunes)
could be thought as the middle stage． It showed that
the third stage of succession process was the most sta-
ble community． In order to prevent degeneration of S．
vulgaris community，the most important thing to do is
to protect the third stage．
3 Conclusion
The autogenic succession process of vegetation in
Mu Us Sandland is from P． uillosa + A． arenarium
(A． sphaerocephala)on semi-drift-sand dunes to Ar-
temisia ordosica + S． vulgaris on semi-fixed-sand
dunes，then through S． vulgaris + P． sphondylodes
on fixed-sand dunes and S． vulgaris on fixed dunes of
subclimax stage to S． vulgaris + moss community on
old-fixed-sand dunes，which maybe pass to a S． bun-
geana steppe finally． Within the succession process，
the coverage of community and species diversity al-
ways changed． The species diversity increased with
the development of the progressive succession，it was
the most abundant in the third stage，and then de-
creased． From the early stage to subclimax one，the
ecological environment began getting wet． Along with
the declining of S． vulgaris，some plants exited，the
habitat of community began getting dry． The most sta-
ble community is the third stage in the process of S．
vulgaris community． So，the third stage of succession
should be protected in order to prevent the degenera-
tion of S． vulgaris community．
Ｒeferences
1． Zhang G S，Dong Z，Wang L H． The researches review of the
bioecology characteristic and the growth-reproduction of Sa-
bina vulgaris［J］． Journal of Inner Mongolia Forestry Col-
lege，1997，19(2) :69 － 75．
(下转 128 页)
3111 期 红雨等:臭柏群落在不同演替阶段某些群落特征的研究
说明大斑病长蠕孢 002(菌 B)与 001(菌 A)在相同
诱导浓度下前者的作用强度相对较弱。
3 株长蠕孢属真菌侵染幼苗期毛脉酸模试验
结果表明:大斑长蠕孢 001 以 a浓度处理毛脉酸模
种子可以对幼苗期毛脉酸模中的二苯乙烯类成分
的含量和产量累积产生显著性影响。其中对白藜
芦醇的影响最明显。大斑长蠕孢 001 对幼苗期毛
脉酸模中白藜芦醇苷产量促进最强时期，处理组是
空白对照组的 2． 54 倍;对毛脉酸模中白藜芦醇产
量促进最强时期，处理组是空白对照组的 4． 99 倍。
Sparvoli等已从葡萄中分离出 Ｒes(白藜芦醇)
生物合成途径中的相关酶基因［11］。党尉等研究认
为 Ｒes生物合成途径需要一系列相关酶的协同作
用。当加入诱导因子后首先相关酶的 mＲNA 含量
有所上升，随后蛋白质水平也上调，因此证明的确
合成了部分酶［12］;另外也有可能是植物体内原有
的相关酶活性较低或处于钝化状态，诱导物则作为
酶的激活剂发挥作用。也许正是这种双重作用使
得诱导可以明显提高 Ｒes的含量。
本实验研究了 3 种真菌对幼苗期毛脉酸模中
白藜芦醇苷和白藜芦醇的诱导规律，经综合分析
后，筛选出了大斑病长蠕孢 001 对幼苗期毛脉酸模
中的白藜芦醇苷和白藜芦醇的诱导效果较明显。
但其作用机制是否符合上述机理有待进一步的研
究。本项研究为下一步继续探寻这种真菌的最适
处理浓度及验证最适处理时间提供了科学依据和
实践指导。下一步将从分子生物学水平进行诱导
途径研究，探讨产生诱导效应的诱导物及其在毛脉
酸模上的作用靶点。继续研究长蠕孢属真菌其他侵
染途径对毛脉酸模中的白藜芦醇苷和白藜芦醇及
其它次生代谢产物的影响。
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821 植 物 研 究 28 卷