全 文 ：热带亚热带植物学报 2005，13(2)：95—104
Jouma／ofTropicol and Subtropical Botany
广东亚热带森林木本植物幼苗生长特性研究
彭姣凤，陈章和 ，韦明思
(华南师范大学生命科学学院，广州 510631)
摘要：研究了广东亚热带42种木本植物幼苗的生长及其与物种的生态特性、生活型、种子大小的相互关系。较强光下
(H，66．8 I~mol m-2s )乔木幼苗的茎高和茎生物量显著高于灌木幼苗的相应值，但在较低光．卜(L，33．7 I~mol m-2s- )两
者无显著差异。而阳性植物、耐阴植物和中间型植物之间，茎高和茎生物量无显著差别。乔木幼苗的叶面积和叶生物量
比灌木幼苗大，但灌木幼苗的叶片数较乔木幼苗多。大种子种和小种子种幼苗之间，阳性植物、耐阴植物和中间型植物
幼苗之间的叶片数和叶面积一般无显著差异。阳性植物幼苗比耐阴植物幼苗侧根数多。乔木幼苗的根生物量和根 ／茎
比显著高于灌木幼苗。在较高光下，阳性植物幼苗的根 ／茎比较耐阴植物幼苗高，但在较低光下无明显差异。45 d幼苗
的根生物量与种子重量呈显著的正相关，而90 d幼苗无明显的相关。乔木幼苗个体生物量显著高于灌木幼苗。幼苗相
对生长率和叶面积比的大小呈现如下顺序：阳性植物>中间型植物>耐阴植物，但只有阳性植物和耐阴植物之间有显
著差异。阳性植物、中间型植物和耐阴植物幼苗之间的单位叶率无显著差异。
关键词：幼苗生长：木本植物：亚热带森林：广东
中图分类号：Q948．1 文献标识码：A 文章编号：1005—3395(2005)02—0095—10
Characteristics of the Seedling Growth of W oody Plants
in the Subtropical Forests in Guangdong Province
PENG J i ao—feng， CHEN Zhang—he’，WE I M i ng—s i
(Colege ofLife Science，South China Normal University，Guangzhou 510631，China)
Abstract：Seedling growth of 42 woody species of subtropical forests in Guangdong Province was studied in rela—
tion to their ecological characteristics， growth forms and seed weight． Seedlings of tree species had significantly
higher stem and greater stem biomass than those of shrub species under high light(H，66．8 I~mol m s )，but not low
light intensity(L，33．7 I~mol m s )．There was no signifcant diference among heliophilic species,shade—tolerant
species and intermediate light—demanding species in stem height and stem biomass．Tree seedlings had more leaf
area and leaf biomass per plant than shrub seedlings，but the latter had more leaves．Diference in leaf number and
leaf area was generally not obvious between large—seeded and small—seeded species，and betw een diferent types of
species based on light demand．Seedlings of heliophilic species had more lateral roots than those of shade—tolerant
species．Seedlings of tree species had signifcantly greater root biomass and higher root／shoot ratio of biomass
than those of shrub species．Seedlings of heliophilic species had also higher root／shoot ratio than those of
shade—tolerant species under H as compared with L condition．Root biomass was significantly positively correlated
with seed weight for 45 d seedlings in both H and L light,but not for 90 d seedlings．Very signifcant linear core—
lation between seedling biomass and seed weight was found for 45 d seedlings under both H and L light
conditions，but not signifcant for 90 d seedlings under H light condition．Tree seedlings had signifcantly greater
biomass per plant than shrub seedlings．Relative growth rate and leaf area ratio of seedlings were foun d to be in
Received：2004-04-09 Accepted：2004-09-14
Foundation item：Supported by the National Natural Science Foundation ofChina(No．39270140，39670138)
Corresponding author
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热带亚热带植物学报 第 13卷
order of heliophilc species>intermediate light—demanding species>shade—tolerant species，and there were signi_
ficant di ：rence between heliophilic species and shade—tolerant species．Difference in unit leaf rate was not signi—
ficant among species ofvarious light—demanding types．
Key words：Seedling growth；Woody plants；Subtropical forest；Guangdong Province
Inhabitation and growth of seedlings are very
susceptible to environmental variation．Seedlings have
to cope with the efects of favorable and unfavorable
environmental factors in their habitat during their
growt h．Therefore，studies on characteristics ofseedling
growt h are of importance for reforestation and for
research of forest dynamics． This study is carried out
based on the folowing considerations：First，although
studies on this aspect have led to an encouraging
understanding on the ecological adaptability to their
environm ent of the species， researches have been
focused mostly on the species in tropical forests[ ]and
temperate forests[~3]，not much research work has been
carried out in subtropical humid forests[ 4， ．Have the
seedlings the same adaptive strategies as those in
tropical and temperate forests? If not。what is the
diference between them?Second，studies have been
carried out mostly on the comparison ofa few species
with obviously different ecological characteristics，such
as pioneers and late—successional species[ ．However．
species might present continuous vari~ion in the
characters of adaptation to a certain environm ent[S,t6, ．
How does the ecological adaptation of seedlings vary
continuously?This paper studied 42 woody species of
diferent light demands or successional phases and of
diferent growth form s as well as seed weights in the
subtropical forest in China． in order to examine these
questions which are important for un derstanding the
ecological adaptation patern of woody plant seedlings
in a certain environmental gradient．
1 Materials and Methods
Seeds of 42 woody species(appendix 1)were
collected during July l 999 to August 2000 from
Dinghushan Nature Reserve,Heishiding Nature Reserve,
Luofushan Nature Reserve，and Baiyunshan Forest Park
in Guangdong Province，which are located at similar
latitudes(23。08’-23。22’N1 with similar climatic con—
ditions：annual average temperature is about 20—22℃．
the coldest(January)and the hotest month(July)tem—
peratures are 12—14℃ and 27—28~C，respectively．Th e
mean annual rainfall is about 1 700一l 900 I／IlTI，the wet
season is from April to September and the dry season
from October to March．The species studied were
divided into shade—tolerant species，heliophilic species
and intermediate species according to their light
demands which were estimated by their appearing fre—
quency inside or outside the forests； into tree species
and shrub species according to growth form； and into
large—seeded species(seed weight>25 mg)and smal-
seeded species(seed weight<7 mg、according to their
seed weight(appendix l 1．Seeds or fruits were mostly
collected directly from mother plants， and in some
cases， freshly shedding seeds or fruits were collected
on the groun d around the mother trees．They were
held in herm etic plastic bags，and then taken back to
the laboratory．Fresh weights were measured by
weighing 50 seeds with electronic balance(mode：
FA1 104，Shanghai)，except for the species with very
tiny seeds such as the F／cus species， such tiny seeds
were measured by thousand seed weight．Dry weight
was obtained after the seeds were dried in an oven
(mode：SKG一01·，Huangshi)for48h at 80~C．
Seeds were germ inated and seedlings were grown
in growth room with day and night temperatures of
25±lcc and 20~1~C，respectively，a relative humidity
ofabout 80％，and day／night hours of 12／12．Seedlings
with similar height were transplanted into plastic pots
．(diameter 12 cm，height 12 cm)filed with cleaned
sand with 1 plant for each pot，50-60 individuals for
each species．PoRed plants were put in suficient space
to avoid interference from each other and competition
for light They were randomly divided into two groups，
one under high light(H，66．8 Ixmol m s )and the
other under low light intensity(L，33．7 Ixmol m s )of
40 W mercury fluorescent lamps and 1 00 W tungsten
filament lamps．Each seedling was alternately supplied
with 1 0 ml Rofison nutrient solution 0 ]one day and
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第2期 彭姣凤等：广东亚热带森林木本植物幼苗生长特性研究 97
l 0 ml deionized water the next day during the first 45 d
and then the volume of nutrient solution and water was
raised to 1 5 ml thereafter． Seedlings were harvested
after 45 d and 90 d of transplanting．Generally，each
time 1 0-1 5 seedlings for each species grown at each
light level were harvested and examined for shoot
height，root length，root number，leafnumber and leaf
area(the later was measured with Delta·T Area Mea·
sure System，England)．Fresh weights of leaf,stem and
root of each seedling were measured．Dry weight of
the organs was measured A1 1 04 electric balance,
ShanghaO after drying for 48 h at 80~C．
Analysis of variance(ANOVA)and a two—sample
t-test were made for the significance of the diferences
for diferent species groups[19J．For abnormal data，a
transition byN／(1+x)to norm al distribution was made
before ANOVA． Correlation analysis was made
betw een tw o variances[~．
2 Results and Analysis
2．1 Growth of seedlings
2．1．1 Stem growth
Stem growth varied in species with difierent
growth forms．Shrub seedlings had lower height and
stem biomass than tree seedlings， and the difference
was signifcant in treatment with hi gh light intensity
l8
l5
l2
9
6
3
O
300
25O
20o
l5O
lOO
so
O
Shrubs
(H)(Fig．1 A，D)．In consideration of that tree species in
general have larger seeds than shrub species【20,2”，
corelation analyses were made respectively for tree
species and shrub species between seed weight and
stem height， and betw een seed weight and stem
biomass． The results showed that no sign ificant
corelations were found between seed weight and stem
height both in tree and shrub species．Corelation
betw een seed weight and stem biomass was sign ificant
for tree species，but not sign ificant for shrub species．
These indicated that sign ificant diferences betw een
the growt h form s in stem height and stem biomass
were in some extent independent of seed weight．Tree
seedlings had significant biomass increment from 45 d
tO 90 d．
There was no sign ifcant diference in stem
height and biomass between the types of light demand，
although seedlings of heliophilic plants had faster
height growth and more biomass increment than those
of shade—tolerant plants， and intermediate light—
demanding plants had medium values(Fig．1 B，E)．
Seedlings of diferent types responded diferently to
light intensities．The heliophilic species grew taller and
had greater stem biomass in H than in L light，while the
shade—tolerant species had higher values un der L than
under H condition．Biomass increment from 45 d to
90 d of heliophilic seedlings was significantly faster
Shade- I眦ei、media~e ]teliopb{li
toleIltillt
口45d 1t 90d f{ 豳,15d L 口 90d L
Sjndl1．
seeded
Large
seede({
Fig．1 Comparison ofseedling stem height and stem biomass among diferent types ofwoody species in the subtropical forests
—H=0一 l【| 【|兰 一 兰一 ≈兰0【l一暑 J|
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热带亚热带植物学报 第 13卷
in I：I than in L．and no significant diference between
the tw o light conditions was found for shade—tolerant
species．Intermediate light-demanding species showed
transitional characteristics between heliophilic and
shade—tolerant plants(Fig．1B，E)．
Stem height and stem biomass were positively
correlated with seed weight in diferent species，how—
ever，the efect of seed weight on stem height and stem
biomass was less for 90 d seedlings than that for 45 d
ones(Fig．1 F，Fig．2)．This indicated that seedlings of
smal—seeded species had higher stem growth rate．
2．1．2 Leafgrowt h
Tree seedlings had fewer leaves but more leaf
area and greater leaf biomass than shrub seedlings
(Table 1 1．These diferences were afected by light
intensity and varied with seedling age．Tree seedlings
from 45 d to 90 d had significant increase in leafnum-
ber，leaf area and leaf weight，and had more leaf area
and leaf weight under H light than L light condition．
90 d shrub seedlings did not have significantly more
leaf number and leaf area but had significantly greater
leaf weight under both L and H light conditions as
一 5 0 5 10 —5 0 5 10
S0o(1 wcig}1t(IIg，10g)
Fig．2 Correlation between seed weight and stem height of45 d and 90 d
seedlings under low(L)and high(H)light intensities
Table 1 Leaf number，leaf ale~ and leaf weight of seedlings
Sh，shade tolerant species；In，intermediate light demanding species；He
，
heliophilic species；Sm，small—seeded species；La
，
large—seeded species；H，high light intensity；L’low light intensity．Numbers folowed by the same letter within the column are not
signifcan tly diferent at p<0．05 for small leter
，and at p<0．01 f0r capital leter．The same for Tables 2 an d 3．
5 0 5 0
5 0 5 0
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第2期 彭姣凤等：广东亚热带森林木本植物幼苗生长特性研究
compared to 45 d seedlings． Shrub seedlings under H
condition had more leaf weight increment than under
L condition．
There was no significant correlation in scatter
diagrams of leaf number，leaf area and leaf weight
against the species in the order from shade—tolerant
species to heliophilic species(the ord．er is aranged in
appendix)；and there was no signifcant diference in
leaf num ber，leaf area and leaf weight among the
heliophilic species， shade—tolerant species and inter—
mediate light—demanding species(Table 1)．However，
leaf growth responded diferently to light intensities．
Heliophilic species had more leaf num ber and greater
leafweight under H than those under L condition，and
had significantly more leaf area and greater leaf
weight in seedlings transplanted after 90 d than after
45 d． Leaf num ber of the shade—tolerant plants was
similar under the two light conditions，whereas leaf
areaandleafweightwerehigherunderLthan underH
condition． Th ere was no sign ificant diference in leaf
area and leafweight between 45 d and 90 d seedlings
of the shade—tolerant plants．These facts showed that H
light condition was more favorable for leaf growth of
heliophilic seedlings than L light condition，and that
shade—tolerant plants needed less light for leaf growth
than heliophilic plants．Seed weight had no signifcant
efect on leafnum ber．Diference in leafweight betw een
large-·and small-seeded species was sign ificant for
45一day seedlings in both H and L light，and diference
in leafarea was foun d only for 45一day seedlings in L
light(Table 11．
0．S
0．4
0．3
0．2
0．1
0
1．2
1．0
0．8
0．6
0，4
0．2
0
2．1．3 Root growth
Root biomass was related to growth form ofthe
species．Shrub seedlings had lower root biomass per
plant than tree seedlings． Seedlings of heliophilic
species had sign ificantly more lateral roots than those
of shade-tolerant species； they also had larger incre-
ment of root biomass from 45 d to 90 d seedlings an—
der both H and L lights．There was generaly no signi—
ficant diference in length ofmain root and root weight
betw een heliophilic species and shade—tolerant specie~
Correlation analysis indicated that root biomass was
very signifcantly corelated with seed weight for 45 d
seedlings in both H and L light，but not signifcant for
90 d seedlings in neither H nor L light(Fig．3)．Difer—
ences in length ofmain root and root biomass between
the large-seeded species and the small-seeded species
were sign ifcant in the 45 d seedlings，but not sign if—
cant in the 90 d seedlings． These indicated that seed
weight had more important efect on root growth at
early stage of the seedlings，and showed less efect
with the development of the plants．
2．2 Biomass and its allocation
2．2．1 Total biomass
Total biomass showed great variation among
species．Under H light,90 d—seedlings ofspecies such as
Broussonetia papyrifera,Podocarpus fleuryi，Sterculia
Zanceolata,Ficus hispida and Ligustrum sinense had
biomass more than 1 200 mg， but the values of
which in Ilex triflora and Saurauia tristyla were less
than 40 mg．Under L light，such species as Ormosia
0．4
0．3
0．2
Q．1
0
0．8
0，S
0 4
0．3
0，2
0．1
0
Q．0t Q．t l t0 tOO lO∞ O
． 1 l tO tO0 tO00
S¨ d_tight 钿g，log)
Fig．3 Correlation between seed weight and root biomass ofseedlings
^w H_-童々 _1 口奇
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1oo 热带亚热带植物学报 第 13卷
pachycarpc~Broussonetiapapyriferc~Podocarpusfleuryi
and Sterc 2 lanceolata had total biomass more than
l 000 mg，and Cratoxylum ligustrinum and Saurauia
tristyla,lessthan 50mg．
The tree seedlings had significantly higher
biomass per plant than the shrub seedlings rFig．4A)．
Correlation analysis respectively for the tree and shrub
species demonstrated that there was sign ificant COrC-
lation between seed weight and total biomass in tree
seedlings(except seedlings of90 d in H lighO，but not
inshrub seedlings．Diferenceintotalbiomassbetween
large—·and small·-seeded species was sign ificant for
45 d seedlings，but not for 90 d ones(Fig．4C)．These
suggested that the diference in total biomass betw een
tree and shrub seedlings was indeed related to growth
forms of the species，although the efect of seed
weight might involved during the early period of
seedling growth．Tree seedlings had very significant
biomass increment from 45 d to 90 d under H light，
but the shrub species had no sign ificant diference in
biomass between 45 d and 90 d seedlings un der both
HandLlight conditions．
Th ere was no significant corelatio
．
n in scater
diagrams of total biomass per plant against species in
the order from shade-tolerant species to heliophilic
species．However，diferences were foun d in response
to light intensity among heliophilic species，shade-
tolerant species and intermediate light·-demanding
species．Shade-tolerant species had higher biomass in
L than in H light，whereas heliophilic species had
higher biomass in H light than in L light rFig．4B)．
Biomass per plant of shade-tolerant seedlings was not
sign ificantly diferentbetween 45 d and 90 d seedlings,
：
．
fI b l
whereas that of heliophilic plants was significant．
Seedlings ofthe intermediate light-demanding species
had a growth rate lying betw een the other two types
(Fig．4B)．
Corelation analysis indicated that biomass per
seedling was positively corelated with seed weight，
but the corelation was not sign ificant for the 90 d
seedlings in H light．Diference in total biomass
betw een seedlings of large-seeded species and those of
small-seeded species was very significant for 45 d
seedlingsbutnotsignifcantfor90dseedlings ig．4C)．
2．2．2 Biomass alocation
Biomass allocation varied among species．Th e
proportion of root biomass comprised more than 30％
ofthetotalbiomassforboth45 dand 90d seedlingsof
Gossampinus malabarica,Sterculia lanceolatc~Brous-
so~tia papyrifero, Ficus lacor and Ficus microcarpa
(90 d seedlings of Gossampinus malabaricum under
L light comprised 64％ of the tota1)， whereas
Psychotria rubr~ A~acia conflea and Ligustrum
sinense， only about 15％． Th e proportion of root
biomass to total biomass and stem biomass to total
biomass increased generally with the growth of
seedlings，but that of leaf biomass to t0ta1 biomass
showed a contrary tendency．
Biomass allocation had relation to growth forms
of the species．Seedlings of trees had higher root／
shoot ratio than those of shrubs．A linear upward
trend was showed in the scatter diagram s of root／shoot
ratio against the species in the order from shade-tolerant
species to heliophilic species， but this trend was not
significant． It was shown also by ANOVA and t-test
for the types of light demanding that significant
Trees Shade Intemediate Heliophilic
tolerant species species
口45d H 困90d H 一45d L 国90d L
Fig．4 Comparison oftotal biomass between diferent types for 45 d and 90 d seedlings under low and high light intensity treatments
O O O O O O ∞ ∞ ∞
一暑Ⅱl一鬻1lo1声葛。工
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第2期 彭姣凤等：广东亚热带森林木本植物幼苗生长特性研究 lOl
diference was found only between heliophilic species
and shade-tolerant species in H light(Table 2、．There
was no significant diference betw een large- and
small-seeded species．
2．3 Growth analysis
2．3．1 Relative growth rate
Th ere was no sign ificant linear trend in scatter
diagrams of relative growth rate(RGR)against the
species in the order from shade-tolerant species to
heliophilic species，although the heliophilic species
had highest RGR，and was in the order of heliophilic
species>intermediate species>shade-tolerant species．
Sign ificant diferences were foun d between heliophilic
species and shade-tolerant species，and between large-
and smal-seeded species(Table 3、．Corelation analysis
showed that there was a significant negative corelation
between RGR and seed weight both in H and L lights．
Th ere was no significant diference between tree
seedlings and shrub seedlings，although the former had
sli曲tly higher values．
2-3．2 Unit leafrate
Most of the species grown under H light had
higher unit leaf rate(ULR)than those under L light，
and a few species(for example Ormosia pachycarpa，a
shade-tolerant forest tree)showed signifcant higher
ULR under L light compared to H light． There were
generally no sign ificant diferences between diferent
growth forms and between diferent types of light
demanding(Table 3、．Corelation analysis between
seed weight and ULR indicated that signifcant
corelation was found in L light but not in H light
(Table 3、．
2-3-3 Leafarea ratio
Leaf area ratio(LAR)was in the same order as in
RGR and ULR for the light-demanding types．There
was sign ificant diference between shade-tolerant and
heliophilic species．No sign ificant diference was
found between tree and shrub seedlings，although the
former showed slightly higher values(Table 3、．
Corelation analysis showed that LAR in seedlings
was very significantly negatively corelated with seed
Table 2 Root／shoot ratio of seedlings for diferent ecological types and life forms
For abbreviations see Table 1
Table 3 Growth analysis of seedlings of species with large or smal seeds,
andwith difere ntlight demands and growth forms
For abbreviations se Table 1
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lO2 热带亚热带植物学报 第 13卷
weight both in H and L lights
3 Discussion
Shrub seedlings have more leaf number，less leaf
area，lower leaf area ratio，and less leaf weight than
tree seedlings and have similar length of main root，
number of lateral roots compared to tree seedlings
although shrub seedlings have remarkably lower root
weight．This indicates that shrubs have no growth
advantage in shoot growt h but advantages to some
extent in root growth as compared to tree specxes．
Generally speaking，shrubs have less growth advantages
in comparison with tree species，which is indicated by
their lower values of shoot height，shoot weight，total
biomass and RGR．Their adaptive advantage might lie
on higher ability to cope with environment stress(such
as deep shade，dry,insect aRack，etc．)．
Diferent plant types of light demand show
diferent responses to light intensity in plant growth．
Heliophilic species show higher values such as leaf
biomass，stem biomass，root biomass，total biomass in
H than in L light， whereas in shade—tolerant species
such values are higher in L than in H light，and the
intermediate light—demanding species having medium
characteristics between them．However，most of the
values measured show no significant diference among
the three light—demanding types．This suggests that，
firstly，even the seedlings of the heliophilic species
need only moderate amount of light m，and diference
in light demands among various light demanding types
is not so great at their seedling stage as at sapling and
adult stages，which has been observed in seedling and
sapling growt h of some species[221；and secondly，there
is a gradient variation in light demanding from
heliophilic species to shade—tolerant species．Th is is
supported by the results that the indices of growt h of
the seedlings measured in this study，such as growth
rate and total biomass,are in the sequence of heliphilic
plants>intermediate light—demanding species>shade—
tolerant species．
Seed weight has obvious efect on root and shoot
growth，and on biomass increment of the seedlings，
especialy at the early stage(45 d)．Fennerl201 reported
that seedlings from large—seeded species had higher
shoot／root ratio for initialy prior capture of light
rather than minerals． But there was no sign ificant
diference in the ratio of shoot／root biomass betw een
seedlings of large—and small—seeded species in this
experiment． However， seedlings of small—seeded
species had larger diference between H light and L
light in leaf growth(including leaf area and leaf
weight)than those of large—seeded species．It is
suggested that one of the efective strategies by which
seedlings from smal—seeded species strengthen their
ability to capture light is to increase leaf area ratio
and not shoot／root ratio． Seedlings from smal—
seeded species have also relatively more lateral roots
and longer main root per unit root weight to achieve
larger root surface area to atain higher growth rate
than large—seeded—seedlings， as has been observed
by Swanborough and Westoby and Wright and
Westoby ．
Result ofthe experiment is showed that there is a
gradient in growth and response to light intensity in
the species．Some values are significantly diferent
only between the heliophilic and shade—tolerant species．
Th is suggests that methodically，integrated screening
program(ISP) for a big lot of species would help
to reveal the gradient variation of plants and that
comparative studies on ecologically distinct species
would help to probe the diferences betw een them．
Both the research procedures could complement each
other in revealing the relationship betw een diferences
in environment and species perform ance．
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热带亚热带植物学报 第 13卷
Continued
Shr,shrub：Tr,tree；Sh，shade—tolerant species；In，intermediate light demanding species；He，heliophilic species；S，smal。seeded species；
L，large-seeded species．
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