全 文 ：武汉植物学研究 2001, 19( 5) : 369～376
Journal of Wuhan Botanical Research
沙地云杉构件动态与结构

邹春静

　韩士杰　徐文铎
(中国科学院沈阳应用生态研究所陆地生态过程开放研究实验室, 沈阳　110016)
摘　要: 研究了沙地云杉的构件种群。结果表明: ( 1)沙地云杉根系生长迅速; ( 2)针叶寿命随
树龄而变化; ( 3)枝条数量与枝条年龄存在如下关系: B40= 2 682. 5e- 0. 695 8A , B200=
5 807. 5e- 1. 311 2A ; ( 4)雄花/雌花随树龄迅速上升; ( 5)不同层次和方向上球果数量存在显
著差异; ( 6)东方向的冠半径最长, 西方向最短,枝条数量、长度,小枝数量随树龄而增加,小枝
长度随树龄而缩短。结果表明,沙地云杉能够适应干旱的生态环境,其构件动态与结构是长期
的自然选择与适应的结果。
关键词: 沙地云杉; 构件; 动态; 构筑型; 沙地森林; 过渡带
中图分类号: Q944; Q948. 11　文献标识码: A　文章编号: 1000-470X ( 2001) 05-0369-08
Modular Dynamics and Structure of Picea mongolica
ZHOU Chun-Jing
**
, HAN Shi-Jie, XU Wen-Duo
(L abor atory of T err est rial Ecological P rocess, Inst itut e of Ap pl ied Ecology , The Chinese
A cademy of Sc iences, Sh enyang　110016, China)
Abstract: The modular populat ion o f Picea mongolica is studied in this paper.
The results show ed that : ( 1) Root o f Picea mongol ica grow s rapidest in order
fo r adapt ing the drought environment ; ( 2) Needle life changes w ith the t ree
age, needle life o f 40 year -o ld t ree is longer that of 200 year-old t ree; ( 3) As to
br anches, there are r elat ionships between branches number and branches age,
tho se ar e: B40= 2 682. 5e- 0. 695 8A ( R
2= 0. 955 3) , B200= 5 807. 5e- 1. 311 2A
( R
2
= 0. 960 3) ; ( 4) T he rat io of male f low ers and female f low ers increases
rapidly with t ree age; ( 5) T her e is g reat signif icance of cones r at io in dif ferent
layers and direct ions with age. T he t ree is o lder, the r at io is larg er in m iddle
layer, and the rat io is few er in upper lay er . The ratio is larg er in southern and
w estern direct ions, and it is few er in northern and eastern direct ions; ( 6) As far
as crow n type and architecture type is concerned, the radius o f crow n in easter n




Author for corresponden ce.E -mail : chjzou@ iae. ac. cn
Received date: 2001-01-09, Accep ted date: 2001-06-20。
Found at ion items : The Nat ional Natural Science Foundation of Ch ina( No. 39670133, 39900019, and 30070129) .
　Biograph y: Zou Chun jing ( 1968- ) ,m ale, Ph. D. , as sociate profes sor.
direct ion is longest , that in western direct ion is sho rtest . Branches number,
br anches leng th, small branches number, and branches ang le increase w ith t ree
age, but leng th of small branches decr eases. T he results show ed that Picea
mongolica could adapt to the drought environment . M odular dynam ics and
st ructure of P icea mongolica result f rom long-term natural select ion and adap-
tat ion to the drought environment .
Key words : Picea mongolica; M odule; Dynam ics; Architectur e; Sandy forest ;
Ecotone
In sandy fo rest-steppe ecotone, there ex ists a special sandy fo rest ecosystem—Picea
mongol ica forest , w hich has play ed an important role in ameliorat ion of local peoples life
and control to desert if icat ion. P icea mong olica is an endem ic and endangered species in
China
[ 1]
. For Picea mongol ica, populat ion dynamics is v ital eco logical pro cess for it s ex is-
tence and extension
[ 2]
. How ever , w e must re-understand plant indiv idual in some as-
pects. Higher plant is a modular or ganism, w hich is continually replicated by the incr ease
of basic modular units such as leaves, buds, branches, tillers and so on. M odular o rga-
nisms ar e ex tr emely flex ible in size, form , number and g row th rate of modular units in
response to their internal and ex ternal condit ions. Perennial w oody plants ar e architec-
tures formed by modular units most of w hich are dead t issues [ 3] .
The populat ion-like st ructure of a higher plant is emphasized by the w ay in which
plants are classif ied. Almost the whole of the info rmat ion used by taxonom ists in the de-
script ion of plants involv es features o f the leaves, f low ers, seeds, et al., that is the modu-
lar units of const ruct ion. T he fo rm o f the w hole is tax onomically almost irrelev ant be-
cause it is so enormously variable. T he rel iable repeatable unit is the constr uct ional mo-
dule, not the w hole. In this paper, we studied modular dynam ics and str ucture of Picea
mongol ica for expr essing its adaptability , and hoped to provide scient ific basis fo r conser-
vat ion of the endangered t ree species in sandy forest-steppe eco tone[ 4] .
1　Site description and methods
1. 1　Site description
The study w as conducted in Baiyinaobao Natural Reserve, at the eastern area of In-
ner M ongol ia Autonomous Region, China, w ith 43°30′～ 43°36′N lat itude, 117°06′～
117°16′E long itude, and 1 300～1 500 meters alt itude. The Reserve locates at edge o f
Hunshandake Desert , the t ransit ional region from Da Hinggan Ling Mountains to Mon-
golia Plateau. T he highest mount is Aobao Mount w ith 1 498. 8 meters alt itude. T he hill
is composed of volcanic lapil lus and basalt , and other places are all covered by sand. T he
sand layer r ange is 10～100 meters thick [ 5] .
Climate here is typical temperate cont inent steppe climate. Annual mean tempera-
ture is - 1. 4℃, and mean temper ature is - 23. 4℃ in January and 14. 7℃ in July. M ean
370 武汉 植 物学 研究　　　　　　　　　　　　　　　　　第 19卷　
annual rainfall is 448. 9 millimeters w ith only one rainy season in summer f rom June to
August , and taking up 68% of that o f the w hole year. T he potent ial evapo ration is 1 526
millimeters. Zonal soils o f the ar ea are mainly black so il and br ow n soil, w hich distr ibute
on dunes.
1. 2　Methods
Seedl ings modular grow th and dynamics. 2～5 seeds w ere sow ed in 3 cm×3 cm pat-
tern on April 9, 1995, and they germinated and became seedlings af ter 15 day s. We sur-
veyed seedlings height , main root leng th, small roo t length, needle length, stem height ,
biomass of root , needle, and stem in each month.
M odular characterist ic of individuals. W e adopted life table to survey needle number
and age st ructur e. Numerical characterist ic of br anches, reproduct ive org ans w ere sur-
veyed. Some standard tr ees o f P icea mongolica wer e selected, and divided the canopy into
thr ee layers. We selected some standard branches in each lay er , and surveyed cone pro-
duct ion o f Picea mongolica in eastern, southern, w estern, and nor thern dir ect ions, and
analy zed them in statistical methods.
Architectur e type o f P icea mongolica. Canopy radius, branches number, branches
leng th, smal l branches number, small branches length, and branches ang le, and so on
w er e surveyed.
2　Results
The plant g row th is the format ion of it s modules over and over again. T herefo re, we
could consider the modules to be individuals, and take the w hole plant as a modular po-
pulat ion.
2. 1　Modular growth and dynamics of Picea mongol ica seedl ings
Seeds o f Picea mongolica germinate and g row into seedling s af ter 15 days, and then
enter a rapid-g row th stag e ( T able 1) .
Table 1　Modules growth of Picea mongolica seedlings
Dt
Da
( d)
H (cm) Mrl ( cm ) S rn L n ( cm) H s (cm)
Wd ( g)
Ro S t Ne
Sn
5. 18 39 6. 38±0. 717 2. 84±0. 696 - 1. 11±0. 129 2. 31±0. 209 0. 106 9 0. 161 7 0. 319 1 100
6. 20 72 9. 83±2. 053 6. 26±2. 007 - 1. 14±1. 076 3. 57±0. 480 0. 597 1 0. 316 1 0. 842 3 100
7. 19 102 14. 81±2. 919 10. 75±2. 928 4. 04±1. 955 1. 15±0. 135 3. 65±0. 689 1. 274 2 0. 359 9 1. 000 2 100
8. 20 134 16. 53±2. 950 13. 05±3. 108 4. 10±2. 100 1. 16±0. 212 3. 99±0. 704 1. 542 1 0. 497 5 1. 107 5 100
9. 19 164 18. 48±3. 133 14. 33±3. 329 4. 53±1. 821 1. 18±0. 135 4. 10±0. 383 2. 065 6 0. 517 6 1. 197 6 100
10. 20 194 22. 35±3. 625 18. 36±3. 658 4. 67±1. 660 1. 18±0. 104 4. 18±0. 850 2. 294 0 0. 520 6 1. 415 2 100
Notes : Dt - date; Da- days; H - heig ht ; Mr l- main root length; Srn- sm all root num ber ; Ln- needle length;
Hs- stem heigh t ;Wd- dry w eigh t; S n- sample number; Ro- root ; St- stem; Ne- n eedle.
　　We analyzed the modules rat io of P icea mongolica seedlings in or der for expressing
their modules dynam ics ( T able 2) .
371　第 5期　　　　　　　　　　　　邹春静等:沙地云杉构件动态与结构(英)
Table 2　Ratio of modules of Picea mongolica seedlings
Dt Da( d) Ha/ Hu S t / Ne Wr/ Wn Wu/ Wa Wn / Ws
5. 18 39 0. 802 3 2. 081 1 0. 335 0 0. 2223 1. 973 4
6. 20 72 1. 752 5 3. 131 6 0. 685 1 0. 4982 2. 664 7
7. 19 102 2. 647 8 3. 173 9 1. 273 9 0. 9368 2. 779 1
8. 20 134 3. 750 0 3. 439 7 1. 392 4 0. 9608 2. 226 1
9. 19 164 3. 453 0 3. 474 6 1. 724 8 1. 2043 2. 313 8
10. 20 194 4. 601 5 3. 542 4 1. 621 0 1. 1850 2. 718 4
Notes: Ha- Height above gr ou nd; Hu- height un der ground; St-
s tem; Ne- needle; Wr - root weight ; Wn- needle w eight ;
Wu- w eig ht u nder ground ; Wa - w eigh t above ground;
Ws- s tem w eigh t.
　　 During g row th process, roo t
gr ow s quickest no mat ter roo t
leng th and roo t biomass. Roo t
leng th increases from 0. 8 t imes of
stem leng th to 4 t imes, and roo t
w eight increases f rom 0. 2 times of
stem and needle to 1. 2 times. Nee-
dle grow s slow ly, but stem g row s
rapidly. As to biomass accumula-
tion, needle biomass is bigger than that of stem .
We surv eyed branches height and their biomass of a f iv e-year-o ld seedling, and the
results are as fol low s ( T able 3) .
Table 3　Height and biomass of branches of 5-year-old seedlings
As 1s t 2nd 3rd 4th 5th
H( cm) 4. 96±0. 911 3. 39±0. 882 2. 43±0. 620 2. 45±0. 560 2. 45±0. 460
Hs( cm ) 3. 38 4. 81 3. 19 2. 62 2. 67
Wn( g ) 23. 78 12. 04 3. 69 0. 31 0 　
　　Note: A s- ag e scale.
　　T here is no sig-
nif icant difference in
stem height and stem
biomass. How ever ,
needle biomass of
one-year -o ld is bigg est .
2. 2　Modular characteristic of Picea mongolica adult individuals
Needle, branches, f low ers, cones and so on are basic modules of Picea mongolica,
and their number and characterist ic w ill direct ly influence physiolog ical pr opert ies and
adaptability of Picea mongolica.
2. 2. 1　Age structure and distribution of needle
The modular str ucture of plants g ives them an age st ructure. Picea mongol ica, like
most clonal plants, has a clearly defined modular st ructure in w hich the genet develops
as a populat ion of modules. For example, needle has it s own dynamics. N eedle passes
thr ough phases o f juvenility to maturity and senescence. The needle starts as a depen-
dent st ructure dr aw ing on resources f rom o ther parts of the plant-it matures and be-
comes itself a net exporter of org anic materials, reaching a peak of act ivity and then de-
clining . Photosynthet ic ef ficient and transpirat ion intensity have relat ionship with needle
age. T herefo re, we established the life table of needle of 40 year -old and 200 year -o ld
t rees ( T able 4 and Table 5) .
Table 4　Needle lif e table of a 40-year-old
Picea mongolica individual
X Ix Dx Qx Lx T x E x
0 1 000 131 0. 131 935 3 405 3. 405
1 869 100 0. 115 819 2 470 2. 842
2 769 111 0. 144 714 1 651 2. 147
3 658 50 0. 076 633 937 1. 424
4 608 608 1. 000 304 304 0. 500
Table 5　Needle life table of a 200-year-old
Picea mongolica individual
X Ix Dx Qx Lx T x E x
0 1 000 272 0. 272 864 1282 1. 282
1 728 674 0. 926 391 418 0. 574
2 54 54 1. 000 27 27 0. 500
372 武汉 植 物学 研究　　　　　　　　　　　　　　　　　第 19卷　
　　In the l ife table, there are many items, and their signif icances ar e the follow ing [ 6] :
X - age scale; Ix - surv iv al individuals in beginning o f X period ( normalization) ;
Dx- the dead individuals f rom X to X+ 1 period; Qx- the dead rat io f rom X to X+ 1 pe-
riod, and that is Dx / Ix ; Lx - surv iv al individuals during X and X+ 1 per iod, and it is
〔Ix+ I( x+ 1)〕/ 2; T x- individuals fr om X to more than X per iod; Ex - mean expected
life of survival individual f rom beg inning of this age scale, and it is T x / Ix .
As far as 40-year-old individual is concerned, mortality of needle is small, and mean
expected life is big . As to 200-year-old individual, mor tality is big, and mean expected
life is very small. T he reason is that the tr anspirat ion rate increases rapidly w ith age in-
creasing, and the cast needle could save w ater to adapt to drought climate. We draw nee-
dle dist ribut ion of these tw o tr ees ( F ig . 1) .
Fig. 1　Distribution of Picea mongolica needle
Needle life of 40-year -o ld tr ee is longer than that of 200-year-old tr ee. M ost needle
distr ibute in m iddle and down part of canopy . In upper part , the needle is few , especially
fo r o ld needle, w hich has relat ionship w ith branches dist ribut ion and canopy form .
2. 2. 2　Numerical dynamics of branches
We took the ment ioned tw o trees as example. T heir branches and branches age are
as fo llow s ( F ig . 2) .
Fig. 2　Branches age structure ( norm al izat ion)
Br anches number and br anches age
could be expr essed w ith the follow ing for-
mula:
B40= 2 682. 5e- 0. 695 8A ( R
2= 0. 955 3) ,
B 200= 5 807. 5e- 1. 311 2A ( R 2= 0. 960 3) 。
Wher e B is branches number, A is
branches age.
With age incr easing , br anches fal l o ff
seriously . We found that tw o-year old needle
and branches fall o ff in sunny slope of dunes, which is a normal r esponse of P icea mon-
golica to drought cl imate.
373　第 5期　　　　　　　　　　　　邹春静等:沙地云杉构件动态与结构(英)
2. 2. 3　Numerical characteristic of reproductive organs
Table 6　Reproductive organs and their
ratio of Picea mongolica
As 31～50 51～70 71～90 91～150 151～250 251～
Sn 9 5 7 5 9 4
Ff 63 302 590 793 903 891
Mf 3 250 21 511 53 395 95 636 143487 150 757
M/ F 51. 6 71. 2 90. 5 120. 5 150. 9 169. 2
Notes : Ff- female f low ers; M f- male f low ers;
M / F- m ale flowers / female f low ers .
In early spring , 1995, w e surveyed
the repr oduct ive organs ( male f low ers
and female f low er s ) and their r at io
( T able 6) .
With t ree age increasing, female
flow er s incr ease gradually . How ever,
male f low ers increase rapidly . There-
fo re, the rat io of male f low ers / female
flow er s incr eases rapidly .
2. 2. 4　Numerical variation of cones
The tw o fundamental components of fitness of a populat ion are repr oduct ion and
survival . By invest igat ing cone product ion of Picea mongol ica in differ ent layer and dif-
ferent direct ions, w e found that cone product ion in upper lay er is more than that in mid-
dle layer and down layer , and cone product ion in south and w est is mo re than that in
north and east .
Picea mongolica beg ins to product from more than 20 years old. W ith the age in-
creasing, seed production incr eases ( T able 7) .
Table 7　Cone production of Picea mongolica
Sn
D
( cm)
H( m) As
Ld
S W N E
Lm
S W N E
L u
S W N E
4 0～5 2. 6～3. 7 22～32 0. 25 1. 00 0. 75 0. 25 2. 50 1. 75 1. 50 7. 25 7. 25 6. 00 5. 25 5. 50
12 5～10 3. 5～6. 9 30～52 1. 10 1. 05 0. 80 2. 10 9. 00 7. 20 9. 00 10. 65 19. 00 20. 75 22. 25 19. 67
23 11～20 4. 2～9. 5 39～85 4. 00 5. 10 4. 67 5. 50 16. 50 16. 75 20. 50 18. 75 34. 39 34. 13 26. 91 30. 83
11 21～30 7. 8～11. 0 70～170 5. 82 5. 73 5. 27 6. 73 36. 18 36. 28 27. 91 40. 73 84. 82 83. 00 69. 55 75. 00
9 31～40 9. 1～14. 0 90～220 11. 11 9. 78 7. 33 8. 78 53. 33 40. 78 45. 77 43. 56 106. 78 98. 67 87. 56 94. 32
11 41～50 11. 0～15. 0 95～240 24. 00 17. 22 19. 56 25. 66 124. 00 96. 78 106. 00 106. 67 157. 89 147. 44 100. 44 128. 44
7 51～　 13. 0～23. 0 150～350 19. 33 23. 67 17. 33 11. 67 294. 33 208. 00 93. 00 163. 00 381. 00 258. 67 196. 00 207. 67
No tes: D- diamet er; Ld- down layer ; Lm- m iddle layer ; L u- upper lay er; S- southern direct ion; W- west ern direct ion;
N- northern direct ion; E- east ern direct ion.
Table 8　Ratio of cone production ( % )
D( cm) Ld Lm Lu S W N E
0～5 6. 47 24. 46 69. 06 28. 78 25. 18 21. 58 24. 46
6～10 4. 12 29. 25 66. 63 23. 74 23. 66 26. 15 26. 45
11～20 8. 84 33. 25 57. 91 25. 18 25. 68 23. 89 25. 26
21～30 5. 15 26. 51 68. 33 27. 77 27. 35 22. 47 26. 49
31～40 6. 09 30. 18 63. 73 28. 17 24. 55 23. 14 24. 13
41～50 8. 20 41. 12 50. 68 29. 02 24. 80 21. 44 24. 74
51～ 3. 84 40. 47 55. 68 37. 07 26. 17 16. 35 20. 41
Notes : D- diameter; Ld- dow n layer; Lm- middle layer; Lu-
upper layer; S - southern d irection ; W- w es tern direc-
t ion; N- northern d irection ;E - eastern direct ion.
Based on this table, w e analy zed
the ratio o f cone pr oduct ion in dif fe-
rent lay er s, differ ent direct ions o f
various age indiv iduals o f Picea mon-
golica ( Table 8) .
In this table, we could see that
there is great significance of cones
product ion rat io in dif ferent layers
and dir ect ions w ith age increasing .
374 武汉 植 物学 研究　　　　　　　　　　　　　　　　　第 19卷　
The tree is o lder, the r at io is larger in m iddle lay er , and the rat io is few er in upper layer.
The r at io is lar ger in southern and w ester n direct ions because of intensiv e light , and the
rat io is few er in northern and eastern directions.
2. 3　Architecture type of Picea mongolica
Architectur e type of a species, w hich is cal led modular st ructure, r efer s to spat ial ar-
rangement o f plant modules. As to Picea mongolica, branches number, branches length,
small branches number, smal l branches leng th, branches angle, canopy st ructure and so
on decided architecture type of P icea mongolica ( T able 9, T able 10 and Table 11) .
Table 9　Canopy range of Picea mongolica
As ( a) 51～100 101～150 151～200 201～
Sn 5 10 9 7
Dr( cm ) 20. 4～35. 5 36. 4～53. 2 46. 1～65. 2 59. 2～75. 0
Dm ( cm) 27. 1 41. 9 54. 1 63. 4
Hr ( m) 7. 5～11. 5 10. 0～18. 0 16. 0～21. 0 17. 0～23. 5
Hm(m ) 9. 3 13. 4 18. 9 20. 5
Rc E 2. 6 4. 0 4. 8 5. 3
( m ) S 2. 6 3. 6 4. 4 4. 6
W 2. 0 2. 8 3. 4 4. 2
N 2. 7 3. 9 4. 6 4. 7
Notes : Dr- diam eter range; Dm- diameter mean
value; Hr- high ran ge; Hm- high m ean
value; Rc- can op y radius .
Table 10　Architecture type of Picea mongolica
As 7～50 51～100 101～150 151～200 201～
Sn 23 14 11 9 13
Dr( cm ) 5. 0～18. 0 17. 5～36. 0 32. 5～55. 1 43. 5～66. 4 59. 2～78. 0
Dm ( cm) 10. 9 25. 8 40. 7 53. 9 64. 5
Hr ( m) 1. 3～6. 0 6. 4～11. 9 10. 1～18. 3 17. 0～22. 5 19. 0～24. 0
Hm(m ) 4. 9 9. 5 14. 2 18. 9 21. 2
Nb 33 62 103 209 342
Lb( m) 1. 4 1. 7 2. 0 2. 5 2. 7
Nsb 5 9 13 24 33
Lsb( m) 0. 35 0. 33 0. 28 0. 25 0. 24
Ab 72. 2 78. 3 85. 2 88. 7 89. 8
Notes : Nb- branches num ber ; Lb- b ran ch len gth ; Nsb-
sm al l branches number; Lsb- small branch length;
Ab- branches angle.
Table 11　Canopy structure in dif f erent layers of trees
No .
A
( a)
H
(m )
Ha
( m)
1st
NL L SN SL Ab
2nd
NL L SN SL Ab
3rd
NL L SN SL Ab
4th
NL L SN SL Ab
5th
NL L SN SL Ab
1 12 1. 2 0. 24 - - - - - 5 0. 14 11 0. 09 60 5 0. 28 43 0. 08 64 10 0. 39 105 0. 08 65 17 0. 57 213 0. 08 70
2 16 2. 4 0. 48 9 0. 11 17 0. 08 56 5 0. 28 45 0. 08 61 6 0. 40 84 0. 09 65 9 0. 79 208 0. 08 68 11 0. 95 484 0. 08 73
3 26 3. 5 0. 70 3 0. 12 7 0. 06 65 10 0. 37 71 0. 08 68 8 0. 69 202 0. 11 68 27 1. 01 891 0. 09 69 33 1. 03 1324 0. 08 75
Notes : No- t ree n umber; A- age; Ha- h eigh t of each layer; NL- branches n umber; L- branch length ( m) ;
SN - small branch es num ber ; S L- small b ranch len gth (M ) ; Ab- branch es angle.
Based on the table, eastern canopy radius of P icea mongolica is longest , and w ester n
canopy radius is shortest . T her e is no significant difference in southern canopy radius
and nor thern canopy radius. The reasons may be as follows. T he f ir st reason is that w est
w ind is main w ind in Baiyinaobao Natural Reserve, and the second is that the solar radia-
tion is v ery intensiv e, which limits ex tension of branches to w ester n direction. T his is the
result of long-term natural select ion.
As far as crow n type and architecture type is concer ned, the radius of crow n in eas-
ter n direct ion is longest , that in western direction is shor test . Number of branches,
leng th of branches, number o f small branches and angle of branches increase with t ree
age, but length of small branches decreases, w hich show s top g row th superiority de-
creases w ith age increasing.
375　第 5期　　　　　　　　　　　　邹春静等:沙地云杉构件动态与结构(英)
3　Discussion
The plant individual is a basic unit in the plant eco logy [ 7] . For the past 20～30
years, plant ecolog ists have g radually accumulated new information on the plant indivi-
dual, w hich are discussed as follow s. Higher plant is sedentary. Higher plant is genet ic
mosaic. Higher plant is a modular organism
[ 8] . T he “r amet”is the unit of clonal growth,
the module that may of ten follow an independent existence if severed f rom the parent
plant . In many higher plants, clonal g row th occurs by the informat ion of a populat ion o f
parts of w hat w as orig inally one seedling. Although the unit modules of a single genet
w ill be genetical identit ies this does no t mean that they w il l compete w ith each o ther on
equal term s. Parts of a clone or shoo ts on a t ree w il l dif fer phenotypically, for example,
in age and size, and may of ten make quite different demands on environmental resources.
Pheno typic dif ferentiat ion betw een neighbouring parts of a clone is of ten w ell developed
as ring pat terns in w hich the central ramets in a dense clone are depauperate compared
w ith those at an invading perimeter
[ 6, 7]
. In recent y ears, many eco logists show ed their r e-
search inter ests in plant modular st ructure and dynamics
[ 7 10] . Sum ida did resear ch on
thr ee-dimensional str ucture o f tr ees and forest in mixed broad-leaved fo rest
[ 9] . Paul et
al . established a mathemat ical model on t ree basal area growth and studied the gr ow th
situat ion of loblo lly pine ( Pinus taeda)
[ 10]
. Fr om 1990s, w e did research on P icea mon-
golica forest under support by National Natural Science Foundat ion of China
[ 1, 2, 4]
and
found that P icea mongol ica can adapt to the sem i-ar id sandy env ironment by modifying
its anatomical and phy siolo gical st ructure. M odular dynamics and st ructure ar e critical in
adaptability of Picea mongolica.
References:
[ 1 ]　Xu W D (徐文铎) , Li W D (李维典) , Zh eng Y (郑沅) . A study on taxonomy of P icea mong oli ca in Inner Mon-
g ol ia. B ul l Bot Res, 1994, 14( 1) : 59 68.
[ 2 ]　Zou C J (邹春静) ,Han S J (韩士杰) , Xu W D (徐文铎) . Ecolog ical field theor y applicat ion to regenerat ion of
Picea mongoli ca popu lat ion. A cta E col Sinica, 1999, 19( 1) : 50 54.
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