全 文 ：植 物 学 报
Acta Botanica Sinica 2003, 45 (10): 1210-1217
Received: 2003-01-08 Accepted: 2003-06-08
Supported by the State Key Basic Research and Development Plan of China (G2000018607), the National Natural Science Foundation of China
(39860024, 30070138), the Knowledge Innovation Program of The Chinese Academy of Sciences (KSCX1-08-02), National Science Foundation
of China for Distinguished Youth Scholars (39825106) and Combating Desertification Research Project (FS 2000-009).
* Author for correspondence. E-mail: < dongming@95777.com >.
http://www.chineseplantscience.com
Phenotypic Plasticity in Response to the Heterogeneous Water Supply
in the Rhizomatous Grass Species, Calamagrostis epigejos
in the Mu Us Sandy Land of China
ZHANG Cheng-Yi 1, 2, 3, U Fei-Hai 1, CHEN Yu-Fu 4, DONG Ming 1*
(1. Laboratory of Quantitative Vegetation Ecology, Institute of Botany, The Chinese Academy of Sciences, Beijing 100093, China;
2. National Climate Center, China Meteorological Administration, B ijing 100081, Ch a;
3. Agricultural University of Inner Mongolia, Hohhot 010019, Ch na;
4. Institute of Geographical Sciences and Natural Resources Research, The Chinese Academy of Sciences, Beijing 100101, China)
Abstract: Calamagrostis epigejos (L.) Roth. i a perennial grass with slender and long rhizome segments
between interconnected neighbor ramets. To investigate the phenotypic plasticity in response to the
heterogeneous soil water supply, ramet pairs of the species were subjected to heterogeneous water
supply by which either mother ramets or daughter ramets were in high or low soil water supply, respectively,
in the Maowusu (Mu Us) Sandy Land of Nei Mongol. The results showed that the phenotypic characteris-
tics of the individual ramets of C. epigejos were greatly influenced by the heterogeneous water supply. The
ramets treated with high water supply significantly produced more new rhizomes and more offspring (ramets),
and accumulated more shoot biomass, and allocated more biomass to their shoots than those treated with
low water supply. In comparison with the daughter ramets in homogeneous soil water supply, phenotypic
characteristics, in terms of new rhizome growth, the production of new offspring, and the biomass alloca-
tion pattern, of the daughter ramets within the pairs of the species were not significantly changed, no
matter that high or low soil water supply to mother ramets. The phenotypic responses of mother ramets
to soil water supply were similar to those of daughter ramets. From these results, it is inferred that the
interconnected ramets of C. epig jos r sponse phenotypically to their local soil water rather than to the
soil water experienced by the interconnected ramets. The interconnected ramets of C. epigejos might be
independent of each other in water relationship, although they are physically interconnected with rhizome
segments. The physiological independence of interconnected ramets might facilitate the risk spreading
and thus enhance the genet survivorship under the frequent drought stresses in Mu Us Sandland.
Key words: phenotypic plasticity; heterogeneous soil water; Calamagrostis epigejos ; ramet pair ; rhizome;
clonal plants; Maowusu (Mu Us) Sandy Land
The potential independence of ramets is a distinct fea-
ture of clonal plants (St. Pierre and Wright, 1972; Ashmun
et al., 1982). The physical connection between ramets does
not always allow the clonal integration to occur (Schmid
and Bazzaz, 1987; Price et al., 1992). The clonal integration
inevitably involves some cost, such as energy consump-
tion (Pitelka and Ashmun, 1985). In the habitat with a low
competition and patchy resources, the interconnected
ramets intend to be independent of each other under the
frequent stresses of environment (Pitelka and Ashmun,
1985). For example, reproducing vigor of the shoots of
Gaylussacia baccata, a fire-tolerant clonal shrub, was di-
minished after fire by connection to other shoots with rhi-
zome segments, due to competing for resources stored in
the rhizomes (Matlack, 1997). The sister ramets of Aster
acuminatus, a rhizomatous forest herb, was independent of
each other in the patchily distributed light environment un-
der tree canopy (Ashmun et al., 1982). The newly produced
ramets of Leymus secalinus were independent of their pa-
rental ramets in their establishment (Dong, 1999). In the harsh
environment, the strategy of spreading the ramets into the
much more sites within the habitat and then surviving inde-
pendently of each other by ramets may be a successful
mechanism for the clonal plants to overcome the environ-
mental harshness. The independence of the ramets of clonal
plants may facilitate the risk of the genet mortality to be
spread among its all ramets to gain the evolutional advan-
tage for the genet (Cook, 1985; Dong, 1996). Based on the
subsequent analysis of clonal plants adaptation to their
environments, Cook (1985) argued that the programmed
1211ZHANG Cheng-Yi et al.: Phenotypic Plasticity in Response to the Heterogeneous Water Supply in Calamagrostis epigejos
independence of intraclonal ramets would greatly increase
the probability of the genet survival in the severe
environment. Although the probability of the ramet mortal-
ity was high in the severe environment, the probability of
genet extinction is the product of these separate chances
of death and it would decline with the multiplication of
ramet death (Cook, 1985; Dong, 1996).
In the habitat of the arid and semi-arid inland dunes,
environmental factors, such as dry climate, exposure by
wind erosion, burial by sand accretion, poor edaphic
conditions, impose strong selective pressures on plants
(Zhu and Bao, 1993; Zhang, 1994). The knowledge of the
adaptation of plants to the dune environment can make the
revegetation of the desertified areas practicable.
C. epigejos, growing in the arid and semi-arid inland
dunes of northwestern part of China (Yang, 1994; Cui, 1996),
spreads its ramets into many micro-sites in the habitats by
clonal growth with long and slender rhizomatous segments.
In order to understand the adaptation of the species to the
sand dunes, an experiment of the heterogeneous water sup-
ply to the ramets in pairs of the species was conducted. In
this paper, the results of the experiment on the phenotypic
plasticity of the ramet pairs in response to the heteroge-
neous water supply are reported to provide some funda-
mental knowledge for the selection of plants in combating
desertification.
1 Materials and Methods
1.1 Species and methods
Calamagrostis epigejos (L.) Roth., a perennial rhizoma-
tous grass (Gramineae), distributing in the open habitats in
the temperate zone in the Eurasia (Lu, 1987), often grows
on the habitats with ample water supply such as field margin,
field ridge, river bank and lowland (Xu, 1999). The most
individual ramets of the species grow singly with vertical
stems and with belowground rhizomes in sympodial branch-
ing pattern (Institute of Botany of The Chinese Academy
of Sciences, 1976). The rhizome segments, mainly with 10-
100 cm in length, are persistent and lack of thickening with
the size increase after the ramets’ establishment.
The experiment was carried out in Ordos Sandy Land
Ecological Station of the Institute of Botany, The Chinese
Academy of Sciences. Ordos Sandy Land Ecological Sta-
tion (39°29N; 110°11E) is located in the southeast of Ordos
Plateau of Nei Mongol, with altitude 1 300-1 400 m, annual
mean temperature 6.0-8.5 °C, mean annual precipitation
370 mm. The dunes in this station were dominated by some
shrub species, such as Artemisia ordosica Krasch.,
Hedysarum laeve Maxim., Salix psammophila C. Wang et
Ch. Y. Yang. The plant materials used in this study was
propagated from an original genet collected from the
dunefield in the station. The plant collection was made in
March 2000 and multiplied clonally in a greenhouse until
the start of the experiment in July 2000.
1.2 Treatments
Mother-daughter ramet pairs, interconnected by a single
sympodial rhizome segment, were separated from excavated
stock plants, and then transplanted into plastic pots (height
20 cm, diameter 20 cm). One mother-daughter ramet pair
was rooted in a pair of pots with each of the interconnected
ramets of a pair in a separate pot. The rhizome segments,
connecting mother and daughter ramets between pair of
pots, were surrounded with moistened sand and were sealed
with polyethylene to prevent water loss. After three weeks
of transplanting, 32 ramet pairs with similar size were se-
lected and imposed upon the designed treatments on July
28, 2000. There were two levels of water supply in this
experiment: high water supply (+) and low water supply
(-). For the high water supply, the water content in the
pots of a pair was maintained at 12%-18% (i.e. 67%-100%
of field capacity) by daily adding 200-500 mL water into
the pots. The 50-150 mm water was added to the other
pots to maintain their soil water content at 3%-8% (i.e.
from wilting point to 45% of field capacity) for low water
supply. The high or low water supply was imposed to
mother-daughter ramets (M-D) in a two-way factorial design,
resulting in four experimental treatments: homogeneously
high water supply to both the mother and daughter (M+D+),
heterogeneously high water supply to the mother (M+D-),
heterogeneously high water supply to the daughter (M-
D+), and homogeneously low water supply to both the
mother and daughter (M-D-) for treatments A, B, C, and
D. The individual treatments were assigned randomly to
each of the four treatments and replicated eight times. All
plants were harvested on October eight of 2000.
1.3 Measurements and data analyses
The number of leaves, leaf length of both the treated
ramets and their new offspring were counted and measured,
respectively. The number of the offspring ramets was
counted as well. After harvest, the number of nodes, and
the total length of new rhizome of the individual treated
ramets were counted and measured, respectively. The
above-ground part, the root, and the rhizome biomass of
the individual treated ramets were weighed after dried at 80
℃ for at least 48 h. The root/shoot ratio, and mean length
of internodes were determined by the corresponding
characteristics.
All data were analyzed with two-way ANOVA (procedure
植物学报 Acta Botanica Sinica Vol.45 No.10 20031212
GLM; SPSS) to check the effect of the water supply level to
the plasticity of the clonal growth of C. epigejos (Sokal and
Rohlf, 1981; Spss Inc., 1997). The posterior comparison for
all data between treatments were carried out with Duncan’s
multiple range test (Du, 1999).
2 Results
2.1 The number of new offspring
The water supply to the mother ramets had a significant
effect on the number of the new offspring of the mother
ramets, but did not on the number of the new offspring of
the daughter ramets (Table 1). The effects of water supply
to the daughter ramets on the number of the new offspring
of both the mother and daughter ramets were not signifi-
cant (Table 1). By Duncan’s multiple range test, the new
offspring of the mother ramets in treatments A and B were
significantly more than those in treatments C and D (Fig.1).
The daughter ramets in treatment A and C produced more
offspring than those did in treatment B and D, but did not
significantly (Fig.1).
2.2 The number of leaves and leaf length
The number of leaves and leaf length of the treated indi-
vidual ramets were not significantly varied among the treat-
ments (P > 0.05; Table 1). However, the number of leaves
and leaf length of new offspring of both mother ramets and
daughter ramets were significantly affected by the water
supplies (P > 0.05; Table 1). Under the high water supply
(treatments A and B), the number of leaves and the leaf
length of the new offspring of mother ramets were signifi-
cantly greater than that they did under the low water sup-
ply (treatments C and D); the total number of leaves and
total leaf length of the mother parts under the high water
supply were also significantly greater than those under the
low water supply (Fig.2). The four characteristics, i.e., the
Table 1 Pvalues of phenotypic characteristics of Calam grostis epigejos a analyzed using two-way ANOVA. The factors are
treatment to mother ramet and treatment to daughter ramet
CharacteristicsTreatment toTreatment to Treatment to mother ramet×
daughter rametmother rametTr atment to daughter ramet
Number of newly produced offspring of mother ramet 0.001 0.300 1.000
Number of newly produced offspring of daughter ramet1.000 0.372 0.550
Number of leaves of mother ramet0.186 0.848 0.701
Number of leaves of daughter ramet0.178 0.721 0.445
Number of leaves of new offspring of mother ramet<0.001 0.619 0.956
Number of leaves of new offspring of daughter ramets1.000 0.001 0.817
Total number of leaves of mother plant part0.003 0.595 0.809
Total number of leaves of daughter plant part0.457 0.001 0.831
Length of leaves of mother ramet0.995 0.256 0.825
Length of leaves of daughter ramet0.434 0.585 0.676
Length of leaves of new offspring of mother ramet<0.001 0.463 0.494
Length of leaves of new offspring of daughter ramet0.854 0.003 0.358
Total length of leaves of mother plant part0.002 0.954 0.498
Total length of leaves of daughter plant part0.809 0.042 0.381
Length of new rhizomes of mother ramet<0.001 0.574 0.634
Length of new rhizomes of daughter ramet0.582 <0.001 0.759
Number of nodes of new rhizomes of mother ramet0.003 0.482 0.905
Number of nodes of new rhizomes of daughter ramet0.622 0.005 0.668
Internode length of new rhizomes of mother ramet0.286 0.673 0.505
Internode length of new rhizomes of daughter ramet0.635 0.322 0.946
Biomass of new rhizomes of mother ramet<0.001 0.857 0.857
Biomass of new rhizomes of daughter ramet0.532 0.001 0.965
Shoot biomass of mother ramet 0.997 0.629 0.920
Shoot biomass of daughter ramet0.810 0.932 0.779
Shoot biomass of new offspring of mother ramet<0.001 0.190 0.879
Shoot biomass of new offspring of daughter ramet0.517 <0.001 0.316
Total shoot biomass of mother plant part<0.001 0.217 0.870
Total shoot biomass of daughter plant part0.694 <0.001 0.257
Total root biomass of mother plant part0.010 0.212 0.509
Total root biomass of daughter plant part0.338 0.091 0.290
Ratio of root/shoot of mother plant part<0.001 0.682 0.238
Ratio of root/shoot of daughter plant part0.966 <0.001 0.404
1213ZHANG Cheng-Yi et al.: Phenotypic Plasticity in Response to the Heterogeneous Water Supply in Calamagrostis epigejos
number of leaves, leaf length of the new offspring of daugh-
ter ramets and the total number of leaves, leaf length of the
daughter parts under the high water supply (treatments A
and C) were significantly greater than those under the low
water supply (treatments B and D) (Fig.2). Therefore, under
the heterogeneous water supply, the ramets under the low
water supply and connected to ramets under the high wa-
ter supply did not increase the number of leaves, leaf length
of their new offspring and the total number of leaves, leaf
length of their plant parts (Fig.2).
2.3 The total length, number of nodes, length of intern-
odes and biomass of the new rhizomes
The total length, the number of nodes of the new rhi-
zomes both of the mother ramets and daughter ramets were
significantly affected by the water supplies to themselves,
but were not by water supplies to the connected ramets in
pair (Table 1). The total length and the number of nodes of
the new rhizomes of the mother ramets under the high wa-
ter supply (treatments A and B) were significantly greater
than those under the low water supply (treatments C and
D) (Fig.3). However, the mean length of internodes of the
Fig.1. The number of new offspring produced by mother ramets
(left panel) and daughter ramets (right panel) of Calamagrostis
epigejos ramet pairs under the four different water supply
treatments. A, B, C and D represent the treatments in which
mother ramet and daughter ramet were subjected to homoge-
neously high water supply with both mother and daughter ramets
in the high water supply (M+D+), heterogeneously water sup-
plies with mother ramet in the high water supply and daughter
ramet in the low water supply (M+D-), heterogeneously water
supplies with mother ramet in the low water supply and daughter
ramet in the high water supply (M-D+), and homogeneously
low water supply with both mother and daughter ramets in the
low water supply (M-D-). The bars in the same panel with the
same letter are not significantly different at P = 0.05.
Fig.2. The number of leaves (1), and leaf lengths (2), produced
by mother ramets and their offspring (left panel), daughter ramets
and their offspring (right panel) of Calamagrostis epigejos ramet
pairs under the four different water supply treatments. A, B, C
and D are described as in Fig.1. Open portions, shaded portions
of bars, and whole bars denote separately the values of the treated
ramets, their offspring, and plant parts; and were labelled with
letters a and b, l and k, x and y, respectively . For the individual
portions or the whole bars of the same figure, values with signifi-
cant difference at P = 0.05 were labelled with different letters.
Fig.3. Length of new rhizomes (1), the number of total nodes of
new rhizomes (2), length of new rhizome internodes (3), and
biomass of new rhizomes (4), produced by mother ramets (left
panel) and daughter ramets (right panel) of Calamagrostis epigejos
ramet pairs under the four different water supply treatments. A,
B, C and D are described as in Fig.1. The bars in the same figure
with the same letter are not significantly different at P = 0.05.
植物学报 Acta Botanica Sinica Vol.45 No.10 20031214
new rhizomes of both the mother ramets and daughter ramets
was not significantly affected by the water supply (Table 1)
among the treatments (Duncan’s multiple range test, P >
0.05) (Fig.3). The effect of the water supplies to the mother
ramets on the biomass of the new rhizomes of the mother
ramets was greatly significant; however this effect on the
biomass of the new rhizomes of the daughter ramets did
not reach the significant level (Table 1; Fig.3). Similarly, the
effects of the water supply to the daughter ramets on their
new rhizomes was significant while that on the new rhi-
zomes of the mother ramets was not (Table 1; Fig.3).
2.4 Aboveground biomass
The biomass of either the mother ramets or daughter
ramets among the treatments was not significantly differ-
ent (Table 1; Fig.4). The effect of the water supplies to the
mother ramets on the aboveground biomass of the new
offspring of the mother ramets was significant, but did not
reach the significant level on the aboveground biomass of
the new offspring of the daughter ramets (Table 1). Similarly,
the water supply to the daughter ramets had a significant
effect on the aboveground biomass of the offspring of the
daughter ramets, but not on that of the mother ramets (Table
1). The effects of both the water supplies to the mother
ramets and to the daughter ramets on either the biomass of
the mother parts or that of the daughter parts were the
same as those on the biomass of the mother ramets and the
daughter ramets, respectively (Table 1). It was shown that
the biomass of the offspring of the mother ramets and that
of the mother parts under the high water supply in treat-
ments A and B were significantly greater than those under
low water supply in treatments C and D; the biomass of the
new offspring of the daughter ramets and that of the daugh-
ter parts were significantly greater in treatments A and C
than those in treatments B and D (Duncan’s multiple range
test, Fig.4). In other words, though the mother ramets in
treatment C connected with the daughter ramets under the
high water supply, its offspring and their parts did not have
signif icantly more b iomass than those under the
homogenously low water supply in treatment D; the biom-
ass of the new offspring of the daughter ramets and that of
the daughter parts in treatment B were similar to those un-
der the homogenously low water supply in treatment D
(Fig.4).
2.5 The biomass of roots
The effect of the water supplies to the mother ramets
was significant on the biomass of the roots of the mother
parts, but did not reach the significant level on that of the
daughter parts; the effect of water supplies to the daughter
ramets was not significant on both the biomass of roots of
the mother parts and that of the daughter parts (Table 1). It
is shown that the biomass of the roots of the mother parts
under the high water supply in treatments A and B was
greater than that under the low water supply in treatments
C and D; the biomass of the roots of the daughter ramets
was not significantly different among the treatments (Fig.
5).
2.6 The root/shoot ratio
Fig.4. Aboveground biomass of mother ramets and their off-
spring (left panel), and daughter ramets and their offspring (right
panel) of Calamagrostis epigejos ramet pairs under the four dif-
ferent water supply treatments. A, B, C and D are described as in
Fig. 1. Open portions, shaded portions of bars, and whole bars
were described and labelled as in Fig. 2. For the individual por-
tions in the same figure, values with significant difference at P =
0.05 were labelled with different letters.
Fig.5. Root biomass of mother ramets and their offspring (left
panel), and daughter ramets and their offspring (right panel) of
Calamagrostis epigejos ramet pairs under the four different wa-
ter supply treatments. A, B, C and D are described as in Fig.1.
The bars in the same panel with the same letter were not signifi-
cantly different at P = 0.05.
The effect of the water supplies to the mother ramets
was significant on the root/shoot ratio of the mother parts,
but was not on that of the daughter parts. The effect of the
water supplies of the daughter ramets was significant on
the root/shoot ratio of the daughter parts; but was not on
that of the mother parts (Table 1). The interaction of the
water supplies to both the mother ramets and the daughter
ramets did not significantly affect the root/shoot ratio of
the mother parts and the daughter parts (Table 1). The root/
shoot ratio of the mother parts under the high water supply
in treatment A and B was around 0.4, which was significant
smaller than that under the low water supply in treatments
C and D (Duncan’s multiple range test, Fig.6). The ratio of
1215ZHANG Cheng-Yi et al.: Phenotypic Plasticity in Response to the Heterogeneous Water Supply in Calamagrostis epigejos
the root/shoot of the daughter parts under the high water
supply in treatments A and C was also significant smaller
than that under the low water supply in treatments B and D
(Fig.6). The root/shoot ratio of the daughter parts in treat-
ment B and that of the mother parts in treatments C were
significantly different from those under the homogenously
high water supply in treatment A, but had no significant
difference from those in the homogenously low water sup-
ply in treatment D (Fig.6).
2000). The natural habit can be easily observed in the field
where the rhizome segments of C. epigejos do not grow
further strong after the establishment of the new offspring
in the new microsites and are persistent of the connection
between mother and daughter ramets in the original slen-
der and long growth form; even some rhizome segments
decay to split the connection between ramets. According
to the phenomena above, it can be inferred that under the
heterogeneous water supply the interconnected ramets do
not share water or only share little water between each
other, resulting in no significant change of phenotypic
characteristics. In other words, the daughter ramets after
their establishment are independent of their mother ramets
for water, and the mother ramets do not gain water subsidy
from their connected daughter ramets.
In Mu Us Sandy Land, where the experiment was
conducted, the severe environmental stresses and distur-
bances (e.g. drought, defoliation by herbivores, and root
exposure after a severe wind-erosion) occur frequently.
Also, the heterogeneous habitats extensively exist in the
Sandland, such as heterogeneous soil water condition
(Zhang et al., 1997). Imposed on these severe stresses and
the heterogeneous habitats, plants might adopt diverse
strategies to survive in this habitat. Whether or not the
interconnected ramets of the clonal plants share resources,
and the pattern of this resource sharing may be individu-
ally specified for each species (Schmid and Bazzaz, 1987).
For H. laeve growing in the Mu Us Sandy Land, its rhizome
segments of the new daughter ramets after establishment
grow more massive and become thick in size as the daugh-
ter ramets grow. The interconnected ramets can share wa-
ter and are physiologically integrated with each other
through this viable rhizome connection, resulting in the
growth of individual ramets remotely responds to the re-
source supply of the microsites of the connected ramets
(Zhang and Dong, 2000). However, the clonal integration
contributed little to the produce and establishment of the
new daughter ramets of L. secalinus, a clonal grass. An-
other clonal grass—Psammochloa villosa responded the
resource heterogeneous supply with an intensive clonal
plasticity (Dong and Alaten, 1999). After the daughter ramets
establishment, the interconnected mother and daughter
ramets of C. epigejos clones individually respond to their
local water supply in terms of growth. This independence
for water might facilitate the genets of the species to in-
crease the survival in the severe drought condition. And
this independence for water might provide an ecological
strategy for this species to sustainable survive in the habi-
tat of the Sandland with the severe drought stress and
Fig.6. Root/shoot ratio of mother ramets and their offspring
(left panel), and daughter ramets and their offspring (right panel)
of Calamagrostis epigejos ramet pairs under the four different
water supply treatments. A, B, C and D are described as in Fig.
1. The bars in the same panel with the same letter were not
significantly different at P = 0.05.
3 Discussion
Under the high water supply, the treated ramets (either
mother or daughter ramets) produced more new rhizomes,
offspring and allocated more biomass to the shoots
(significantly less root/shoot ratio). Furthermore, the newly
produced offspring significantly grew more leaves and ac-
cumulated more aboveground biomass than those under
the low water supply. It is concluded that the growth of
ramets of C. epigejos was greatly affected by the water
supply of soil.
Under the heterogeneous water supply, the growth char-
acteristics of the ramets in the low water supply, although
they connected with the intraclonal ramets in the high wa-
ter supply, was not significantly different from the ramets
in the homogenously low water supply in terms of root
biomass, aboveground biomass and biomass allocation.
The ramets in the high water supply, although they con-
nected with the intraclonal ramets in the low water supply,
had almost the same phenotypic characteristics with the
ramets in the homogenously high water supply. In the rhi-
zome severing experiment, it is shown that the ramets in the
pairs under the heterogeneous water supply had the simi-
lar phenotypic characteristics with the ramets under the
same heterogeneous water supply and with the rhizomes
severed in terms of growth (Zhang’s personal observation,
植物学报 Acta Botanica Sinica Vol.45 No.10 20031216
intensive disturbances. For the practice of combating de-
sertification of the northern China, C. epigejos, together
with other species, might be employed to revegetate some
sandlands with the drought st ress and in tensive
disturbances.
Acknowledgements: We are grateful to Mr. HU Zhi-Zhong,
Ms. CHEN Jin-Zheng, and Mr. LIU Zhi-Mao for their valu-
able assistance in field experiment.
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(Managing editor: HAN Ya-Qin)
1217ZHANG Cheng-Yi et al.: Phenotypic Plasticity in Response to the Heterogeneous Water Supply in Calamagrostis epigejos
中图分类号: Q948.1 文献标识码: A 文章编号: 0577-7496(2003)10-1210-08
收稿日期:2003- 1-08 接受日期:2003- 6-08
基金项目:国家重点基础研究发展规划项目(G2000018607); 国家自然科学基金(39860024; 30070138）; 中国科学院知识创新
工程项目(KSCX1-08-2）; 国家杰出青年科学基金(39825106）; 科技部首都圈(环北京)防沙治沙应急技术研究与示范项目(FS2000-
009)。
*通讯作者。E-mail: 。
(责任编辑:韩亚琴)
毛乌素沙地根茎禾草拂子茅对异质性水分供应的表型可塑性
张称意1,2,3 于飞海1 陈玉福4 董 鸣1*
（1. 中国科学院植物研究所植被数量生态学重点实验室，北京 100093；2．内蒙古农业大学，呼和浩特 010019；
3. 中国气象局国家气候中心，北京 100081； 4. 中国科学院地理科学与资源研究所， 北京 100101）
摘要： 拂子茅(Calamagrostis epigejos (L.) Roth.)为根茎型多年生禾草，具细长根茎。为了探讨拂子茅在异
质性水分环境中的表型差异，在内蒙古鄂尔多斯高原的毛乌素沙地对拂子茅由母株、子株组成的分株对给予了高
水、低水两种不同的异质性土壤水分处理。实验结果表明：土壤水分状况显著地影响着拂子茅分株的生长表型。在
高土壤水分条件下，拂子茅的分株产生的根茎、新生后代分株较多，并使生物量主要分配于地上部分，地上生物量
积累多；在低土壤水分条件下，拂子茅分株产生较少的根茎与新生后代分株，并且分配到根系的生物量明显增大。
在具有一定对比度的异质性土壤水分环境中，拂子茅分株并不因相连的其他分株所处的土壤水分状况而在根茎生
长、新生后代分株的产生和生物量分配等特征上，与同质环境中的具有相同土壤水分状况的分株相比，有明显差
异。这些结果揭示：拂子茅仅以分株的形式对异质性水分供应发生表型反应；相连的克隆分株在向顶向和向基向这
两个基本方向上，不能对另一分株的土壤水分状况在生长表型上发生反应，它们在水分关系上可能是相互相对独立
的。分株的相对独立可能有利于在气候干旱、扰动强烈的沙地环境中实现风险分摊，提高基株的存活几率。
关键词： 表型可塑性；异质性土壤水分；拂子茅；分株对；根茎；克隆植物；毛乌素沙地