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STUDY ON THE INDIVIDUAL GROWTH DYNAMIC OF ADENOPHORA LOBOPHYLLA——THE ROOT SYSTEM GROWTH DYNAMIC IN THE LIFE SPAN

裂叶沙参个体生长动态研究(2):根系的生长动态



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(2)
———根系的生长动态
张文辉1 , 2 祖元刚1
(1.东北林业大学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学系 , 圣卡罗斯 巴西)
摘 要 在裂叶沙参分布区金川县调查了 29 块样地和 1502个不同年龄个体的根
系空间结构和生物量 ,对 5个生长于不同海拔区域的种群进行了比较 ,分析了影响
根系发育的环境因素 。生长于海拔 2700 ~ 2900m 种群根系发育完整 ,主根 、侧根 、二
次侧根 、细根生物量和体积均达到最高值 。高于或低于这一海拔区域 ,种群根系发
育不够充分 。主要原因在低海拔地区是干旱和外界极度干扰;在高海拔地区是低
温。各种群总合的根系生物量(Y , g)与年令(x , 年)关系可以用公式:Y =0.
003783X1.7953 。其他分布于海拔 2300m 到 3400m 的 5个种群生物量(Y , g)与年龄
(x ,年)关系可用一般公式:Y=a.xb表示。
关键词 裂叶沙参种群;根系生长;生物学特征;生物量
STUDYON THE INDIVIDUAL GROWTH DYNAMIC
OF ADENOPHORA LOBOPHYLLA
———THE ROOT SYSTEM GROWTH DYNAMIC IN THE LIFE SPAN
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 , Brazil)
Abstract In the distribut ion area , Jinchuan county , of Adenophora lobophy lla Hong.
population , 29 sample plots were investigated.The root system spatial st ructure and
biomass of 1502 individuals w ith different ages w ere studied , and 5 populat ions at dif-
国家自然科学基金“八五”重大项目资助(39391500)。 收稿日期:1997-3-10
ferent altitudes were compared.The environmental factors which inf luence the root
sy stem development were analyzed too.The root system of the population at altitude of
2700-2900m could develop fully , the biomass and size of taproot , lateral root , sec-
ondary lateral root , thin roo t all reach the maximum.If the habitat is higher o r low er
than this altitude , the roo t systems of the population could not develop well.The most
principal reason for the underdevelopment at the lower alt itude w as assumed to be
drought and external ex treme disturbance , and at the higher altitude w as assumed to be
low temperature.The relation between root sy stem biomass grow th (y , g)and ages
(x , year)could be expressed by the equation:y=0.003783x1.7953 for the whole popula-
tion , and for the other 5 populations f rom alti tude of 2300m to 3400m could be ex-
pressed by the general equation:y =a·xb.
Key words Adenophora lobophy lla populat ion;Root sy stem g row th;Biological char-
acteristics;Biomass
1 Introduction
The roo t system of perennial herb is very important in it s life cy cle , especially for the geo-
phy te w ith the storage root , it is the unique perennial org an , not only accumulates its volume
and improves the sto rage function , but also expands i ts active surface area , improves i ts abso rp-
tive function〔2〕.The development of the root system is direct ly connected with the ability of
adaptation and resistance to the unfavorable envi ronmental factors.At the same time , the root
spat ial st ructure , function and g row th regulation of all life are di rectly related to the grow th and
the decline of population.The research w ork on the st ructure , function of roo t system w as car-
ried out fo r many years.However , the reports on the root system g row th process of perennial
herb and the relation betw een root system g row th and environment are rare , expecially fo r the
endangered perennial herb species.
Adenophora lobophy lla Hong.is a typical endangered plant species〔8〕 , perennial herb.
Recently , as the endangered plant conservation biology att racts mo re and more scientist to pay
more at tentions , A .lobophyl la has been selected as a typical plant to make tho rough study in
China owing to i ts ecological impo rtance.The w orks on genetic diversity and mo rphological
v ariation has been carried out , and achieved a good result〔3-5〕.However , the know ledge on the
root system w as unknown.Undoubtedly , A.lobophy lla as a geophyte with sto rage roo t , the
study on the root system g row th process and spatial structure is a very impo rtant basic w ork for
explaining the endangered process and mechanism.The aims of this research are to reveal the
general root system g row th and the spatial structure developing regulation of A.lobophylla
population;to compare the difference of spatial st ructure and g row th process among 5 popula-
tions at alti tude of 2300-3400m;and to explain the relations between the root system grow th
and principal environmental factors in order to provide the reference for the analysis on dynamic
of the populations.
1291 期            张文辉等:裂叶沙参个体生长动态研究(2)
2 Materials and Methods
In the northwest of Sichuan Province , Jinchuan County , the A .lobophyl la population
distributes f rom altitude 2000m to 3500m〔6〕.
29 sample plots of A .lobophyl la population w ere disposed at different altitudes f rom
2300m to 3400m.Every difference of altitude at least there are about 2 plots.Because the num-
ber of A .lobophyl la individuals is sparse in the field , and it is difficult to dispose the sample
plo ts at random.Therefore , the areas where there are more rational age st ructure and high den-
sity of population w ere selected as a sample plot generally.The area of each plot changes f rom
6m
2
to 115m
2.After the environmental factors and plant community factors w ere investig ated ,
all individuals in the plot w ere dug out as careful as possible in o rder to measure the biomass ex-
act ly in laboratory .Acco rding to altitude and environmental condition , 29 sample plots were di-
vided into 5 populations.Since most of the natural vegetation in this area has destroy ed , the
names of every population have to be given by altitude scope.Different populations are:f rom al-
titude of 2300m to 2500m , 9 plots wi th 351 individuals;f rom 2500m to 2700m , 7 plots wi th
282 individuals;f rom 2700m to 2900m , 6 plots w ith 366 individuals;from 2900m to 3100m , 4
plo ts 292 individuals;and from 3100 to 3400m , 3 plo ts w ith 211 individuals.
In the laborato ry , fo r every sample plot , we did following research w orks:(1)the age of
every individual w as indentified by its bud scar on the corona , and all samples were divided into
different g roups according to their ages;(2)the roots w ere put into oven , maintaining 80℃for
96 hours.The reason for this is that some roots are very big , and not easy to dry up;(3)every
part of root system such as corona , taproo t , lateral root , secondary lateral root and thin root
(diameter(D)<0.2cm)or root hair of individuals were separated and their biomass and size
w as measured respectively by elect ron balance (AE 260)and caliper;(4)the biomass and size
of every part of root system at the same age of the same population were calculated statistically
and take the mean for the index of g row th at dif ferent ages.For every population , the mean
biomass and size w as calculated f rom every sample plot.Usually w hen the age of individual is
over 10 years old , the number of individual are not enough for statistics , the individual number
of 2-3 years stage was merged.Finally , the mean biomass and the size of every part of 1502
individuals root systems in 29 sample plots were calculated statistically.
3 Results and Discussion
3.1 The g row ing process and analysis of the whole population
The root system g row th of the w hole population implies the basic characteristic of A.
lobophyl la population.Since the influence of ci rcumstance condition , the grow th indexes of
o ther populations could be modified or diversit fied a lit tle , how ever , the basic characteristic
could be maintained.
The root of A.lobophyl la population belong s to geophy te w ith taproot system , like a car-
rot , w ith st rong storage funct ion.It is the unique organ which is perennial in the w hole plant
130 植  物  研  究               18 卷
body , and its biomass and volume have been the resul t of accumulat ion since germination of the
seed.
The root grow th begins at the t ime of germination of the seed.In the f irst year , A.lobo-
phyl la does not sprout stem above ground , the most material it assimilated is used for the
g row th of root.The length of root g row s very fast during the first year.From spring to au-
tumn , it s leng th reach 5-7cm.And corresponding to taproot , the lateral roo t and root hair
have been shaped.After that , although the roo t leng th still increase as the age increases , the
speed has gone down.Until 21 years old , the mean leng th reaches 22.006cm (Table 1).That
the root system go through to deep soil in short time is the result f rom long evolution and adap-
tation.Under harsh environmental condi tion , the root of A.lobophy lla reaches the deep soil ,
on one hand it could get w ater and nutrient for maintaining normal life , on the o ther hand , it
could delay the stress of w ater deficiency and avoid disturbing f rom ex ternal facto rs and improve
i ts ability of resistance.
Table 1 The mean values of spatial growth of different parts of the root system of A.
lobophyl la population.For every organ , the mean values are from statistics of
1506 individuals in 29 sample plots
Age Corona Taproot Lateral root S econdary lateral root
Thin root
(<0.2cm)
(y) Length(cm)
D
(cm) Num.
Length
(cm)
D
(cm) Num.
Length
(cm)
D
(cm) Num.
Length
(cm)
D
(cm) Num.
Length
(cm)
1 0.511 0.106 1.000 4.036 0.091 0.771 0.753 0.072 6.599 0.757
2 1.328 0.166 1.044 5.978 0.184 0.831 0.890 0.086 1.390 1.450 0.032 7.301 0.914
3 1.553 0.242 1.100 7.419 0.326 1.398 1.421 0.103 1.453 1.491 0.055 7.884 1.914
4 2.649 0.314 1.105 8.118 0.513 1.420 1.905 0.138 1.589 1.523 0.064 8.112 1.580
5 2.867 0.404 1.100 10.156 0.693 1.501 2.597 0.183 1.697 1.589 0.102 8.196 1.909
6 3.126 0.496 1.108 12.186 0.705 1.652 2.830 0.203 2.001 1.590 0.114 8.222 2.764
7 3.436 0.555 1.108 14.252 0.834 1.959 3.230 0.272 2.126 1.578 0.123 8.494 2.846
8 3.870 0.634 1.110 15.605 0.932 2.475 3.643 0.319 2.346 1.604 0.141 8.535 2.979
9 4.276 0.734 1.111 16.121 1.050 2.602 4.012 0.322 2.412 1.634 0.155 8.606 3.020
12 5.324 0.905 1.118 17.680 1.321 2.669 4.671 0.360 2.699 1.999 0.193 10.622 3.401
15 6.506 1.399 1.125 20.301 1.651 2.848 5.531 0.408 2.594 2.214 0.206 10.678 3.468
18 7.642 1.735 1.195 21.922 2.064 2.860 5.900 0.443 3.532 2.589 0.247 11.117 3.516
21 8.379 1.745 1.195 22.006 2.356 2.943 6.065 0.507 3.500 2.883 0.287 10.667 3.114
23 8.789 1.781 1.205 22.010 2.400 3.043 6.065 0.510 3.653 3.037 0.311 8.000 3.022
25 8.798 1.791 1.205 22.060 2.402 3.069 6.080 0.511 3.666 3.160 0.331 3.000 3.010
Generally , the individual of A.lobophyl la has one taproot , nevertheless , there are more
pebbles in soil , which obst ruct the normal g row th of taproot , the taproo t could divided branches
and form 2-3 taproots.Usually , the ability to form branches will be stronger as it s age in-
1311 期            张文辉等:裂叶沙参个体生长动态研究(2)
creases(Table 1).
A .lobophylla could form lateral root system.During the one year the young lateral root
has been formed.As its age increases , the number , leng th and diameter increase accumulative-
ly , until the age of death , the size and biomass of lateral root sy stem could get highest level.
One individual could has 3.069g lateral roots and its leng th is 6.080cm on the average (Table
1).When the grow th of taproot meet obstacles , the lateral roots develop vigorously .Especially
in the loam soil w ith a lot of pebbles , the number , size , biomass of lateral root could reach high
level , comparing wi th that in the other kinds of soil.
The secondary lateral roots g row on the lateral roots.When the individuals of A.lobo-
phyl la is 2 years old , the size of the secndary lateral roots w ould be large enough to be measured
(Table 1).After that , as its age increases , the number , leng th and diameter accumulate contin-
uously .Until i ts age of death , the biomass and size of the secondary lateral roots reach its maxi-
mum.One individual has 3.666g biomass of secondary lateral roots w hich is 2.160cm long , and
diameter is 0.331cm on the average.
The roo t hairs or thin roots(D<0.2cm)play an important role in the w hole life of A.
lobophyl la individual.It is the principal o rgan to get w ater and nutrient f rom soil.When the
radical forms the taproot system , the root hai r begins to execute its function.As the age in-
creases , the number and biomass of the root hair or thin root goes up continuously.Until 18-
20 years old the biomass and the number reach i ts maximum.The g row th of the roo t hair or the
thin root is coordinated w ith the grow th of lateral root system.However , the biomass ratio of
the root hair or thin root to the biomass of all root sy stem will decrease as the all biomass of root
sy stem increase.
The corona of the root of A.lobophy lla is a t ransition between roo t and stem.And i t is
the hub of asexual propagation in the life of A.lobophy lla individual , From corona , the stems
above , base-leaves and the t ransverse stems g row .At the same time , there are many dormant
buds on it.The grow th of the co rona is coordinated w ith the g row th of all roo t , i ts length and
diameter increase as all the root system g row s.Until the dead age , i ts biomass and size reach
the maximum.One individual usually has one co rona on the top of roo t.Nevertheless , when
the habitat is disturbed seriously by the external factors such as being gnawed by the animals ,
being buried and pressed by pebble , gravel or stone , the co rona could branch off and form 2 or
more coronas.This ability could be stronger as the age increases and sto rage function improves.
According to the statistics of biomass w hich is f rom 1506 individuals in 29 sample plo ts at
altitude of 2300-3400m , the biomass g row th process of all population could be divided into 3
periods.Grom 1 to 3 years old , it is young seedling period , the g row ing speed is slow.From 3
to 18 years old , it is vigo rous grow ing period , the growing speed is fast.From 18 to 25 , it is
decrepit period , the grow ing speed is slowed down (Fig .1〈1〉).
3.2 Comparison and analy sis on root sy stem structure and biomass g row th of dif ferent popula-
tions
On the different altitudes scope , there are different comprehensive environmental condi-
132 植  物  研  究               18 卷
tions.Being influenced by the enviromental factors , the size , number and biomass of the indi-
vidual co rona , taproo t , lateral roo t , secondary lateral root and thin root of different populations
all show their dif ferent characteristics respectively.Generally the principal growing indexes of
A .lobophyl la individual mentioned above , increase as the altitude goes up from 2300m to
2900m , and get the highest level at the altitude of 2700-2900m.After that , these g row ing in-
dexes decrease as the altitude increase further(Table 2 , Fig.2).
Table 2 The mean values of morphological characteristic of the root system of A.lobo-
phylla populations at different altitudes of 2300-3400m.For every organ , the
mean values are from statistics of all individuals in the corresponding plots
Population Corona Taproot Lateral root S econdary lateral root
Thin root
(<0.2cm)
Alt.
(m)
Length
(cm)
D
(cm) Num.
Length
(cm)
D
(cm) Num.
Length
(cm)
D
(cm) Num.
Length
(cm)
D
(cm) Num.
Length
(cm)
2300—3400
(all popul.) 4.78 0.71 1.15 14.71 1.32 2.04 3.84 0.30 2.73 2.06 0.14 8.98 2.45
2300—2500 5.06 0.65 1.03 12.71 1.17 2.08 3.52 0.26 2.33 1.59 0.07 8.79 2.38
2500—2700 4.99 0.73 1.01 14.12 1.36 2.55 3.77 0.29 2.14 1.82 0.10 6.75 2.92
2700—2900 5.10 0.90 1.31 17.89 1.64 2.74 5.09 0.41 3.17 2.27 0.19 9.78 2.70
2900—3100 4.81 0.64 1.08 16.98 1.45 1.26 3.66 0.27 3.16 2.53 0.14 6.16 2.38
3100—3400 4.04 0.53 1.21 11.84 1.01 1.57 3.15 0.30 2.83 2.08 0.10 6.14 2.86
The roo t biomass g row th processes of dif ferent populat ions show individual characteristics
respectively on the given altitude.The biomass grow th patterns of 3 populations at the altitude
of 2300 -2500m , 3500-2700m , 2900-3100m are similar.af ter 3 years old the biomass
g row s fast(Fig.1<2>, <3>, <5>).Although they have the low g row th speed period of
decrepit af ter 18-20 years old , i t is not clear.The biomass g row th pat tern of the population at
altitude of 2700-2900m , like that of the all population could be divided into 3 stages clearly.
From 1 to 3 years old , it is in the low speed g row th stage;f rom 3 to 20 years old it is in the
high speed g row th stage;af ter 20 years , it is in the decrepit period of low speed grow th(Fig.
1<4>).The biomass grow th pattern of the population at altitude of 3100-3400m is particu-
larly different.In the first 5 years , it grow s slow ly;f rom 5 to 10 years old , it grow s rapidly;
af ter that , the biomass almost stops g row th.The another characteristic of this population is
that the life span of root sy stem is only 15 years , about 10 years sho rter than that of other pop-
ulations(Fig.1<6>).
The root sy stem g row th of different populat ions at the given alti tudes is the result of com-
prehensive interact ions between the biological characteristics of populations and environmental
factors.Acclimatizing to different environmental conditions , the populations adjusted their liv-
ing st rategy , and show the dif ferences on morphology , material accumulation and life rhythm
During the w hole life , the roo t sy stem grow th is inf luenced and modif ied by the environmental
1331 期            张文辉等:裂叶沙参个体生长动态研究(2)
Fig.1 The biomass g row th process of different populations from altitude 2300 to 3400m as the age in-
creases.
(1)the biom ass of all population is the statist ical data f rom 1502 individuals in 29 plot s;(2)is from 351 individuals in 9
plot s;(3)is f rom 282 individuals in 6 plot s;(4)is f rom 366 individuals in 7 plot s;(5)is f rom 292 individuals in 4
plot s;(6)is f rom 211 individual in 3 plot s.
factors comprehensively.Especially , the soil conditions such as soil tex ture , soil depth and soil
w ater , play an important role in the root sy stem g row th process.In the loam or loose soil , the
root system develops very w ell , and usually the taproot could get the deep soil , and for deep
root system.The longest could reach 80cm.In the clay or tight soil it could no t develop fully
the lateral roots develop relatively well , and fo rm shallow root system.If in the soil there are a
lot of pebbles or boulders , the taproot is f requently obst ructed , the lateral roots and root hairs
could develop completely.The soil thickness is direct ly in proportion to the development of root
sy stem , especially taproo t.In the drought soil , the root could go down to the deep soil , in o r-
der to obtain more w ater in the soil.However , if w ater is too much in the soil , the root system
could not develop fully.Generally , the organs above ground and root system are integ rated by
the vascular bundle , if the root sy stem grow s badly , the organs above ground also go worse.
Correspondingly , the ability of the individual reproduction and survival sould be weaker , there-
fore , the ex tension ability of all population would be crippled.
At altitude of 2700-2900m , we could assume that there is an optimal composi tion of envi-
ronmental factors fo r A .lobophyl la population to grow.Especially , the temperature is no t so
lower than that at the higher alti tude and the moisture is relative higher than that at the low er
altitude.The disturbance f rom humban being has reduced comparing with that in the lower alti-
tude.These condi tions combining with the vegetation and soil condit ions make the envi ronmen-
tal conditions much better than that in other place.Under this circumstance , the spatial struc-
ture , number disposal , and biomass of root system all reach the optimum.
134 植  物  研  究               18 卷
Fig.2 The comparison of different org an biomass o f 5 populations
The five organs of population at alt.2700-2900m are all signifi cant dif ferent f rom other populat ions;the corona , t ap-
root of population at alt.2900-3100m are significant diff eren t from other populations by t-test (P<0.05).All data
were got in the same w ay as Fig.1
If it is lower than this altitude , although the reasons for the underdevelopment of root sys-
tem are complicated , we could believe that drought envi ronment and the disturbance f rom hu-
man being are the most important factors.In this area most individuals of A.lobophyl la popu-
lation live in the low er part of the slope w here the soil w ater is relative suf ficient , or they live in
the thorny shrub communi ties , avoiding the disturbance f rom animal and human being.
If the alti tude is higher than 2900m , the most principal reason fo r underdevelopment of
root system might be the lower temperature.As the altitude goes up , the rainfall increases , and
drought has no t been restrained factor.The disturbance f rom human being or animals have been
alleviated.The vegetation and soil conditions have been near the natural conditions.The envi-
ronmental factors w hich restriain A .lobophy lla population grow th might be at tributed to low
temperature.The highest limit for A.lobophy lla population grow th is altitude of 3500m.As
for the cause of the short life span of this population , we could assume that the population ad-
justs its st rateg y according to specific environment.However , the exact answer to this question
needs to study further.
The situation above seems that the favorite environmental condition fo r the population of
A .lobophylla is very st raitened.In addi tion to the external factors the A .lobophylla popula-
tion itself has weak abili ty to develop.
3.3 General regulation on the root system of different populations
Although influenced by many envi ronmental factors , the root system grow th processes of
every populat ion show general regularity , especially the biomass g row th.The relations between
the biomass of root system (y , g)and time(x , years)were reg ressed for the different popula-
tions(Table 3).It is clear that the equation:y =a.xb is good for expressing the relation of the
1351 期            张文辉等:裂叶沙参个体生长动态研究(2)
root biomass and ages.
Table 3 The regressive equations of relations between biomass and age of A.lobophylla
populations at different al t.2300-3400m
Altitude of populations
(m) Equations R DF F.Rat F0.01
2300—3400(Whole populat ion) y=0.003783x1.7953 0.98 1 , 13 354.1 9.1
2300—2500 y=0.001773x1.8024 0.96 1 , 10 113.9 10.0
2500—2700 y=0.001703x1.9593 0.96 1 , 14 190.8 8.9
2700—2900 y=0.009935x1.8346 0.93 1 , 10 66.1 10.0
2900—3100 y=0.000528X 2.1444 0.98 1 , 10 343.2 10.0
3100—3400 y=0.000688x1.9274 0.97 1 , 8 153.0 11.3
4 Conclusion
The roo t sy stem grow th of A .lobophyl la population is the result of comprehensive inter-
actions between the biological characteristics and the environmental facto rs.The optimum envi-
ronmental condition is very st raitened , just at alti tude of 2700-2900m .The populations adjust
i ts living st rategy against specific circumstance continuously during all life and form different
g row th pattern.Although root sy stem g row th is inf luenced by many external facto rs during all
life , the basic regulation is manintained.The every part grow th of roo t system is coordinated
w ith one another.The development status of root sy stem is connected wi th the abili ty of popu-
lation ex tension.
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136 植  物  研  究               18 卷
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Shen Xian-sheng et al:Ohserve the Fruits of Lactuca sativa and It s Varieties w ith an Electron M icroscope
P lateⅠ
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沈显生等:图版 Ⅱ Shen Xian-sheng et al:PlateⅡ
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P lateⅠ
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邓彦斌等:图版 Ⅱ Deng Yan-bin et al:PlateⅡ
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张萍等:高山红景天胚胎学研究 图版 Ⅰ
Zhang Ping et al:Embryological S tudies of Rhodiola sachalinensis A.Bor Plate Ⅰ
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张萍等:高山红景天胚胎学研究 图版 Ⅰ
Zhang Ping et al:Embryological S tudies of Rhodiola sachalinensis A.Bor Plate Ⅰ
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