全 文 ：植　　　物　　　研　　　究
BULLETIN OF BOTANICAL RESEARCH
第 18 卷　第 1 期 1998 年　1月
Vol.18　No.1 Jan., 　1998
裂叶沙参个体生长动态研究(1)
———一个生长季地上部分的生长
张文辉1 , 2　祖元刚1
(1.东北林业大学森林植物生态学开放研究实验室 ,哈尔滨　150040)
(2.圣卡罗斯联邦大学植物系 , 圣卡罗斯　巴西)
摘　要　在不同生境条件下的固定样地内 ,观察分析了裂叶沙参种群的地上部分在
一个生长季的生长过程和物候特点 。生长于灌木群落下 ,裂叶沙参地上部分生物量
生长(y , g)与时间(x , d)的关系可以用公式:y =0.2872-0.0187x+0.0009x2 表示;
地上各器官茎 、叶 、花枝 、花芽 、花和果的生物量(y ,克)与生长时间(x ,天)的关系可
以用公式:y=b0+b1x +b2x 2表示。从 4月 10日到 8月 15日的速生期间 ,地上各器
官总生物量(B ,克)与生长时间(A ,天)的关系可用逻辑斯蒂方程:B=0.99/(1+e
(3.613-0.149A))表示。茎和花枝的生物量(B ,克)与生长时间(A ,天)关系可用通
式:B=K/(1+e (c-rm.A))表示 。与生长于开阔草本群落中个体相比 ,生长在落叶
多刺灌丛群落下的裂叶沙参个体生长开始时间(物候)晚 ,结束生长时间早 。
关键词　裂叶沙参;个体生长;物候学;逻辑斯蒂方程;濒危物种
STUDYON THE INDIVIDUAL GROWTH DYNAMIC
OF ADENOPHORA LOBOPHYLLA
———GROWTH DYNAMIC IN ONE GROWING SEASON
Zhang Wen-hui1 ,2　Zu Yuan-gang1
(1.Open Research Labo rato ry o f Fo rest P lant Ecolog y , Northeast Fo restry Univ ersity , Harbin
150040)
(2.Bo tanical Department , Federal University of Sǎ o Carlos , Sǎo Carlos , Brazil)
Abstract　In the fixed sample plot at different habitats , the grow ing process of an en-
dangered species , Adenophora lobophylla Hong population w as observed.The regula-
tion of the adult individual g row th and phenological characteristics was studied.The re-
国家自然科学基金“八五”重大项目资助(No.39391500)。　收稿日期:1997-3-10
lation betw een time (x , day)and the total grow th of biomass above g round (y , g)
could be expressed:y=0.2872-0.0187x+0.0009x 2 , and the relations betw een time
(x , day)and the grow th of biomass(y , g)of every o rgans such as branch , leaf , flow er
branch , flow er bud , f low er and fruit could be expressed by the general equation:y =b0
+b1x+b2x2.During the time of rapid g row th from April 10 to August 15 , the relation
between time(A , day)and the grow th of total biomass of all organs above ground(B.
g)could be expressed by Logistic equation:B=0.99/(1+e (3.613-0.149A));the
relations betw een time (A , day)and the g row th of biomass (B , g)of stem , flow er
branch could be expressed:B=K/(1+e (c-rm.A)).The regulat ion of g row th of in-
dividual in the deciduous thorny shrub community were compared with that in g rass
(drought-resistant)community.The individual in the open grass community begins
i ts g row th(phenology)later and ends i ts grow th earlier than that under shrub commu-
nity.
Key words　Adenophora lobophy lla;Individual g row th phenology;Logistic equation;
Endangered species
1　Introduction
The individual g row th is an impo rtant aspect of plant population ecology.Investigation and
study on the grow th regulat ion of plant individual no t only could explain the basic chain of plant
population life cycle but also could provide the important reference fo r population forecast and
population dynamic analysis.The traditional plant grow th analy sis is concerned with agg regate
quantities like biomass , duration of g row th , relative grow th rate and w ith how these dif fering
between sites or habitat〔2〕.However , up to now , most concerned research w orks concentrate
on one or tw o org ans of plant
〔1〕.The systematical study of the stem , leaf , f lower , f ruit and
root is rare , especially , the work of systematically studying the typical endangered plant species
during one g row ing year.In this research , the long-term phenological observation w as carried
out in field conditions and the g row th in different habitats w as analy zed and discussed.The
aims of this research are to explain the individual grow ing and developing relation of A .lobo-
phyl la population in one year , to analyze grow ing process , and to provide evidence or reference
for the analy sis on the endangered mechanism and endangered process of A.lobophyl la popula-
tion.
2　Materials and Methods
A .lobophy lla is perennial herb〔7〕 , living in the drought -resistant thorny shrub or
drought resistant herb communi ties.Its dist ribution is only on the east boundary of Tibet
plateau , Jinchuan county (the no rth lati tude 31°8′-31°58′, and the east longitude 101°13′-
120 植　　物　　研　　究　　　　　　　　　　　　　　　18 卷
102°19′)in Sichuan province.The total area is not more than 5000km2.It has been known as
a typical endangered plant〔11〕.
Tibet plateau rose rapidly af ter the later Tertiary Period and Quaternary Period , and it has
become the highest plateau in the wo rld.The crust movement and earthquake occurs f requent ly
and formed complex topog raphy.Most places are V -shaped river valley and the altitude
changes f rom 2000m to 5000m〔8 , 10〕.
Below altitude of 2400m , it is warm temperate monsoon climate , w ith drought season
(November-April)and raining season (May -October).The mean annual rainfall is 616.
2mm , and the mean annual temperature is 12.8℃.At altitude of 2400-3000m , it is temper-
ate monsoon climate , At altitude of 3000-4000m , cold temperate climate.
There are a few kinds of soil in this area , which are mainly hilly drab soil , hilly brow n soil
and dark brow n w soil
〔8〕.According to 29 sample plo ts which w e have invest ig ated , the pH val-
ue change f rom 7.1 to 7.8 , the o rganic mater change f rom 2.8% to 7.8%, and the soil mois-
ture changes from 6.6% to 36%.
At alti tude of 2000-2600m , the vegetation is the deciduous thorny shrubs and xero herb
communi ty;at altitude of 2600-3400 , the sub-alpine conifer-broad-leaves forest;at alti-
tude of 3400-4000m , the sub-alpine conifer forest , and at altitude of 4000-4400m , alpine
g rassy marshland.A .lobophy lla lives in those different communities f rom altitude of 2000m to
3500m.
Table 1　The different environmental factors under shrub and grass community on sun
slope(August 10 , 1994)
Habitat
Alti tude
(m)
Coverage(%) Light(lux) Temperature(℃) Moisture(%)
Sh rubs Grass
Soil
surface
In
comm-
uni ty
Soil
(5cm)
S oil
surface
Air
In
commu-
nity
Outside
commu
ni ty
Tanchanggou
in sh rub community
2350 90 30 4390 6550 22.0 25.0 32.5 38 35
Bajiaodiao on
grass community
2500 16 40 4790 15800 27.5 34.5 32.0 36 34
The investigation w as carried out f rom September 1993 to November 1995.4 f ixed sample
plo ts were selected in dif ferent habi tat.Tw o located in the deciduous drought resistant thorny
shrubs in Tanchanggou(alt itude , 2350m).The other two is located in drought resistant herb
communi ty on sun slop in Bajiaodiao (alti tude , 2500m).The reason fo r this is that w e want to
observe the phenological changes influenced by the sun light and temperature(Table 1).There
are 8 adult individuals(8-15 years old)were observed in every plots.The regular interval ob-
servation began at the time of sprout(about April 1), and finished at the time of death of all
parts above g round(about October 30).The contents of observation in the field include mea-
suring the leng th of stem , counting the number of leaves , flowers , f ruit s and spray s.At the
same time , the temperatures of atmosphere , surface soil and deep soil(5cm)were recorded.
1211 期　　　　　　　　　　　张文辉等:裂叶沙参个体生长动态研究(1)
The regular interval time during the vegetative g row th period is 10 days , and during the bloom-
ing and fruit period is 5-7 days.Every g row ing change could not be neglected during this in-
terval time.After every observat ion , near and outside the f ixed plots , 5 individuals w hose age
and size were similar to that of the individuals observed in fixed plots , would be dug out as care-
ful as possible in order to measure the biomass in the laborato ry.
In addition to this , characteristics of seedling , root sy stem and flower & f rui t was ob-
served generally in the field condit ions.
In the laboratory , every o rgan of individual specimen at dif ferent living stage w as separated
respectively as careful as possible and put them in oven(80℃)for 24 hours.Then , the biomass
(weight)of leaves , stems , spray , flow ers , f ruit s of every individual sample were measured by
electron balance(AE 260).At the every growing stage in the same habitat plots , the biomass
of every organ of individual specimen w as calculated statistically more than 6 samples for every
o rgans and go t the mean value as the index of grow th.Finally w e could get the biomass grow th
data w hich is f rom the statistics of every organ of 8 individuals in the same habitat as days in-
crease.In this paper , we mainly discussed the g rowing process of individuals in Tanchangguo ,
and compared w ith that in Bajiaodiaogou.
3　Research Results
3.1　The general g rowing process of the individuals of A.lobophy lla population
The reproduction of A.lobophyl la mainly relies on seed , sometimes on clone.2 cotyle-
dons , go out of soil w hen seed germinate , then turn to green.After germination for about 40
days , the coty ledons w ilt.The seedling of one year old , without stem , just develops 3-7 base
leaves , which are nearly round , few serrate on the margin.When it is more than 2 years old ,
the individual begins to sprout stems and produces flow ers and fruits.As its age increase , the
storing function of root system develops , and the number of stems above ground increase also ,
generally can reach 5-7 stems.If w ater and nutrient in the habitat is enough , the number of
stems could reach more than 20 , which are mo re sturdy and taller , and the number of flowers
and fruits is mo re.However , If the habi tat is barren , it sprouts only 1-2 stems which is thin-
ner and shorter , and the number of f lowers and frui ts is smaller.When the habitat is ex tremely
disturbed by human or the disaster w eather (such as ex t reme drought), it only sprouts base
leaves without stems , When the habi tat is disturbed continuously and seriously , it will propa-
gates by transverse stem (clone).
The root sy stem of A .lobophy lla is perennial.The fleshy taproot system is formed by the
radical af ter seed germination in natural condi tion , and the shape is similar to the carro t〔3-5〕.
During its 1-2 years old , the biomass of roo t system g row s quicker than the stems and leaves ,
When i t is 7-12 years old , the complicate taproot sy stem is formed and the ability of resistance
is stronger.
The corona of the root of A .lobophylla is a t ransition betw een root and stem.However it
is very impo rtant part in the life of A.lobophy lla.From corona , the stems , base-leaves and
the transverse stems g row .At the same time there are many do rmant buds on it.Therefore ,
the corona is the hub of asexual and propagation and g row th.
When A.lobophy lla is more than 2 years old , the individual begin i ts sexual reproduc-
tion.In every spring , the stems(1 or mo re)g row from the corona , and usually there is not
122 植　　物　　研　　究　　　　　　　　　　　　　　　18 卷
branch on the stem .Vegetative g row th goes on for about 3 months , the sing le flower or inf lo-
rescence grow s f rom the leaf axial.The definite panicle inf lo rescence composed of 10-20 small
flow ers , on which the top f low er bloom first.The number of f low er or florescence depend on its
age and habitat.Generally , the number of f low ers increases as its age , and reaches the highest
level(about 70-110 f lowers)when it is 15-21 years old.The bet ter conditions of w ater , nu-
trient , light is , the more the number of flowers is.The number of f low ers on one individual
could reach mo re than 200 in rich habi tat.The period of blooming continues f rom the middle of
July to the end of October (the parts above ground begin to die or wilt).However , for one
small f lower the time of blooming is only for 4 days〔3-5〕.The stamen matures befo re the corona
opening , and disperses the pollens in the first 2 day s of blooming.The pistil matures and re-
ceives pollens in the late 2 day s of blooming.The dominant pollination medium is insects which
are 10 species belonging to 2 families , Apidae and Syphyidae.Four of them , Apis cerana ,
Bombus.sp., Episyrphus bal teatus and Melanostoma scalare , are considered as the efficient
pollinator.
3.2　The g row ing and developing process of every organs above g round in one year
Fig.1　The relation of the biomass grow th of every o rgans above the ground with increasing day s after
sprouting in spring.
(1)In Tanchanggou , under the deciduous thorny sh rubs community , at alt itude 2350m , the grow th period in one
year is 195 days f rom April 1 to October 30;(2)In Bajaodiaogou , in the open grass community on sun slop altitude
2500m , the grow th period in one year is 230 days f rom M arch 16 to November 19.
3.2.1　The stem g row th
The stem begins its height grow th on April 1(the mean atmosphere temperature is 13℃).
After 29 days , the period of its slow g row th , it steps into a stage of the rapid grow th as temper-
ature rises.And this situation continues for 49 days.From July 14(108 days af ter sprouting ,
the mean atmosphere temperature is 20℃), it s vegetative g row th speed is gradually slow down
because reproductive grow th become mo re important.It stops the rapid grow th on August 15.
All org ans above g round begin to w ilt on October 8.Its grow th period in one year is 195 day s
1231 期　　　　　　　　　　　张文辉等:裂叶沙参个体生长动态研究(1)
(Fig.1(1)).The all g row th process of stems could be described by the equation:y=b0+b1x+
b2x
2(y:biomass , g ;x :g row ing days;b0 ,b1 ,b2:parameter)(Table.2).If the equation of the
biomass with time (days)from April 10 to August 15 were regressed , it s g row th regulation
w ould chime in wi th Logistic equation(Table 3).
3.2.2　The leaf g row th
The leaf is the energy source for other organs to grow .However , the leaf g row s on stem ,
and the weight , the area of leaf increases depending on the stem g row s.The all g row th process
could be divided into 4 periods:f rom April 1 to 18 , is the period at low speed , from April 29 to
June 18(29-79 days after sprouting)is the period at higher speed;f rom June 18 to July 14
(79-108 days after sprouting)it s grow th speed is slowed down , and its biomass reach the
highest level on July 14(108 days af ter sprout ing), af ter that , the area and biomass of leaf is
g radually decrease because the leaves on the low er position of stem begin to dies w hile new
leaves increase slow ly or could not increase , until the all parts of individual plant above ground
dies(Fig .1(1)).The leaf g row th regulation chimes in wi th the equation:y =b0+b1 x+b2x2
(Table 2).
Table 2　The regression equations of A.lobophylla population on relation between the
biomass growth (y , g)and time (x , day)in one year (Tanchanggou)under
shrub , at altitude 2350m
Relation Equation
y sum of
square
deviation
Resident
square
deviat ion
Corr.
coef f.
(r)
Total biomass-time(days) y=0.2872-0.0187x+0.0009x2 18.7363 0.0395 0.9989
Stem biomass-time(days) y=0.0994-0.0104x+0.0004x2 2.9665 0.0049 0.9989
Leaf biomass-t ime(days) y=0.0842-0.0042x+0.0004x2 1.1087 0.0150 0.9932
Sp ray biomass-time(days) y=-0.6209-0.0222x-0.0002x2 0.0510 0.0051 0.9484
Flow er biomass-time(days) y=0.2660-0.0635x+0.0013x2 0.0604 0.0249 0.7661
Frui t biomass-t ime(days) y=-22.1186+0.2305x+0.0006x2 0.0281 0.0032 0.9404
3.2.3　The spray (f lower branch)grow th
While the vegetat ive grow th reaches certain level , A .lobophylla begins i ts reproductive
g row th.However , the reproductive grow th starts f rom the spray g row th.From June 2 , the
spray grows f rom axis of leaves (66 days after sprouting).From 67 days to 108 day af ter
sprouting it g row s rapidly (there is a litt le fluctuation).After August 15 it stops g rowing.The
fluctuation of the grow th is assumed drought and rain (Fig.1(1)).This g row th regulation
chimes in wi th the equation:y=b0+b1x +b2x 2(Table 2.)If w e regressed the (spray)biomass
124 植　　物　　研　　究　　　　　　　　　　　　　　　18 卷
g row th w ith the time(days)from June 2 to August 15 , we find that the relation fi ts the Logis-
tic equation(Table 3.).
Table 3 　The Logistic equations of A.lobophylla on relation between stem and spray
biomass growth(B.g)and time A , days from April 10 to August 15
Place Relation
plot.
No. K
rate of
natural
increase
(rm)
Integral
constant
(C)
Logist ic
equation
Corr.
coeff.
(r)
SD
Tanchanggou Days-stem 1 0.99 -0.049 3.61 B= 0.99
1+e(3.61-0.049A) -0.94 0.03
Tanchanggou Days-spray 2 0.47 -0.046 3.67 B= 0.47
1+e(3.67-0.046A) -0.90 0.03
Bajiaodiao Days-stem 16 0.76 -0.041 4.49 B= 0.76
1+e(4.49-0.041A) -0.97 0.05
Bajiaodiao Days-spray 17 0.29 -0.072 9.08 B= 0.29
1+e(9.08-0.072A) -0.87 0.03
3.2.4　The flower g row th
On June 18(82 days af ter sprouting)(Fig .1(1)), the flow er bud appear on the sprays.
The number and biomass of the flow er bud increases as time going on for 47 days.After August
4 it g row s slow ly .After August 25 , few , flowers bud appear in the branch.
The first flower appears on spray on July 14(108 days af ter sprouting).After that , the
number and biomass of flower increase as time going on fo r 20 days.Then , the number of flow-
ers g radually reduces.Until September 8 , Just few flow ers appear on the sprays.The regressed
equation of the biomass of flowers and f low er buds(y , g)w ith time(x , days)is:y=b0+b1x+
b2x
2(Table 2).
3.2.5　The fruit g row th
As the f low er w ithers and falls , the first mature f rui t appears on spray on August 15(140.
days af ter sprouting).On September 8(155 day s), the number of the mature f ruit reach to the
highest level.After that , the number of fruit s reduce rapidly .After October 8(185 days), the
all parts above ground began to wi ther.The biomass of f rui ts(y , g)and time (x , days)could
be regressed to be the equat ion:y =b0+b1x+b2x2(Table.2 , Fig.1(1))
3.2.6　The total biomass above g round g row th
The total biomass above ground include the biomass of leaf , stem , spray , f low er and fruit.
From April 1 to M ay 11 , the to tal biomass maintains low er speed grow th.After May 11 (40
days)the total biomass increase rapidly .From August 4 (129 days)the g row th slow s down
g radually .After August 24(149 days), the total biomass begins to reduce because the leaves ,
flow ers and other org ans wi ther or fall(fig.1(1)).The equation of total biomass g row th (y ,
g)w ith time(x , days)could be expressed by:y=b0+b1x +b2x2(Table 2 , Fig.1(1)).
From April 10 to August 15 the relation of total biomass(B , g)above g round with the time
(A , day s)could be expressed by Logistic equation B=0.99/(1+e (3.613-0.149A)).
3.3　Comparison on the g row th and development of the individuals between under two kind
habitats , under the shrubs and g rass community on sun slop
Comparing the phenological characteristics of the individual under shrubs w ith that in the
g rass community on sun slop , (Fig.1(1),(2)), it is clear that the grow th of A.lobophylla
1251 期　　　　　　　　　　　张文辉等:裂叶沙参个体生长动态研究(1)
that on the sun slop begins earlier 14 day s in spring and finishes later 20 day s in autumn than
that under shrubs , the total g row ing t ime is about 230 days in one year.The period of blooming
of the individual on the sun slop begin at the same time but finishes later about 20 days than
that under shrubs.
The total biomass of individual above ground on the sun slop is less about 25% than that
under the shrubs;the all biomass of leaves is less 9%;the biomass of stem is less 20%;the
biomass of all reproductive organs in less 50%on an average(Fig.1(1),(2)).
On the Logistic equation of the biomass of stem and sprays w ith the time (g row ing)the
environmental capaci ty(K)of the individual under shrubs in the Tanchanggou could reach 0.99
for stem and 0.47 for the spray (Table.3), which are 23% and 38%higher respectively than
those on the sun slop in Bajiaodiaogou.
4　Discussion
Every organs of the plant is integrated by phloem and xy lem.The grow th of all o rgans is
not only regulated by phy toho rmone but also inf luenced by the envi ronmental factors〔9〕.
The biomass of stems and leaves of A.lobophyl la increases proportionally and the biomass
of leaves alw ays is larger than that of stems befo re the time of blooming.From the time of
blooming , the biomass g row th of stem and leaf slows down gradually.When the stem stops it s
g row th(July 14 , af ter sprouting 108 days), the biomass of leaves begins to decrease because
the leaves on the lower position of stem wilt due to the lack of light.Before long , the biomass of
leaves is low er than that of stems(Fig.1(1)(2)).
The grow th of f lowers and fruits is coordinated w ith the g row th of stem and leaf.The veg-
etative g row th is preparation fo r the reproductive grow th.After sufficient vegetative grow ing
and fo rming enough frame fo r f low er and fruit , the spray begins to g row .At the same time ,
the grow th of stems and leaves is slow down or stop in o rder to contribute more energy and sub-
stances to the reproductive organs to produce f rui ts and seeds.The better leaves and stems of in-
dividual grow , the more the individual produces the f lowers and fruits and the better the quality
of seeds are
〔6〕.
Under the arid valley condit ion , the drought alw ay s threatens the g row th and development
of plant , Responding to the fluctuation of the environmental facto rs , the individuals of A.
lobophyl la coo rdinate their size , morphology , number of org ans , and g row th rhythm〔2〕.The
temperature play an important role in the A .lobophy lla phenology.The soil temperature on
the sun slop go up quicker in the spring and go down slower in the autumn than that under
shrub condi tion.The g rowing time of individuals on the sun slop is longer is reasonable.
On the o ther hand , on the sun slop the water def iciency is important for A .lobophylla
life.The individual of A .lobophyl la change their strategy to balance water , such as reducing
size and the number of organs above ground.This is the reason why the every organ biomass of
individual on the sun slop is low er than that under the shrubs.The less vegetative biomass of
stem and leave has to lead the reproductive o rgans to be weaker , further , the all population to
be decline.
Comparing the habitat on the sun slop with that under shrubs it is clear that the habitat
under the shrubs is bet ter.The vegetative and reproductive g row th of individual could develop
completely , and biomass of every organs is larger.The environmental capaci ty (K)in the Lo-
gist ic equat ion of biomass of stems or spray w ith (grow ing)t ime under shrubs is higher than
126 植　　物　　研　　究　　　　　　　　　　　　　　　18 卷
that on the sun slop.On the contrary on the sun slop , the individual alw ays could not develop
w ell , even could not produce flower or f ruit.If most A .lobophylla populations live in this kind
habitat , the population go to be declining is unavoidable.
Unfo rtunately in the no rthw est of Sichuan province the good habitant for A.lobophyla
survive is rare because of the dest ruction from human being .The forest have withdrawn to the
altitude above 2900m , and the most place fo r A .lobophy lla to grow became the desolated arid
g rass slop.The envi ronment condition degrades , soil erodes.All of this show that conservation
of A .lobophylla is an urgent task.The local government should take action to protect the local
environment , stop dest roying the forest immediately and encourage planting t rees and grasses
try to recover environment fo r A .lobophyl la population to survive.
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1271 期　　　　　　　　　　　张文辉等:裂叶沙参个体生长动态研究(1)