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DOI 10.1007/s10265-008-0197-1
REGULAR PAPER
Received: 20 February 2008 / Accepted: 8 October 2008 / Published online: 22 November 2008
The Botanical Society of Japan and Springer 2008
Introduction
Alien plant invasion is a serious threat to natural and seminatural ecosystems worldwide (Mgidia et al. 2007), and
invasive plants that could successfully colonize a broad
range of habitats usually need structural and physiological
acclimation to variable environments and resources (Baker
1974).
Mikania micrantha Kunth and Chromolaena odorata L.
are two of the most notorious invasive weeds in southern
China. Both weeds originate from the Asteraceae family,
with M. micrantha being a perennial herbaceous weed vine
native to tropical Central and South America (Holm et al.
1977a; Cock 1982), whereas C. odorata is a perennial
herbaceous shrub native to Mexico, the West Indies and
tropical South America (Holm et al. 1977b). In its original
region, M. micrantha was abundant in open habitats and
showed poor growth performance under 25% shade (Ipor
1991); C. odorata grew on a wide range of soils and
favored 6070% relative soil water content, but performed
poorly when the relative soil water content exceeded 80%
(Vanderwoude et al. 2005). Although the distribution sites
of the two weeds differ in southern China, they sometimes
occur in the same habitat, such as in Guangzhou. Due to
their high competitiveness, and ability to spread and
establish rapidly, both species have invaded a wide range
of areas in southern China since the late 1980s (Xie et al.
2000; Li et al. 2006), and have caused serious damage to
semi-natural forests, pastures, crops and plantations (Zan
et al. 2000; Feng et al. 2002).
Light, water and nutrients are the main resources regulating the survival, growth and distribution of plants. Most
previous studies on these two species in response to
varying environments each focused on a single factor,
either light or drought (Wen et al. 2000; Wang et al. 2003).
123
70
Site description
123
71
MI
MW
LW
FW
LI
MW
LW
FW
MW
LW
23.6 1.27
13.7 0.40
15.9 0.89 10.1 1.01 23.8 1.42 15.2 0.74 11.8 1.28
36
21.2 1.18
12.6 0.77
10.5 0.56
72
24.4 0.97
16.7 0.45
7.9 0.67
Chromolaena odorata
0
21.9 1.52
14.0 1.45
13.5 0.30 10.6 0.35 26.0 0.83 15.7 1.32 12.7 1.29
36
24.3 1.37
14.1 0.66
11.2 1.20
72
27.1 1.15
18.9 1.87
18.4 0.40
7.3 0.88
498.0 4.90
128.0 2.55
17.2 0.97
36
808.0 16.55
276.0 7.48
19.0 1.18
72
756.0 2.45
256.0 2.45
27.0 1.22
C. odorata
0
490.0 5.16
121.3 1.12
17.8 0.92
36
800.0 12.65
255.0 4.47
20.5 0.45
72
757.5 2.24
257.5 2.24
27.5 1.29
31.5 0.58
31.4 0.57
31.3 0.54
65.9 3.46
70.3 3.10
64.3 3.53
29.8 0.41
29.5 0.39
29.0 0.38
FI Full irradiance, MI medium irradiance, LI low irradiance, FW full water, MW medium water, LW low water
a
Sky conditions: day 0 was cloudy and days 36 and 72 were clear days
Pn
UI Pmax
q
UI Pmax 2 4hUIPmax
2h
Rd
123
72
(a) 20
Statistical analyses
The samples collected were random and all parameters of
leaf physiological and structural characters were used to
test the differences between species and the effects of light
and water treatments on each variable by three-way analysis of variances (ANOVAs) (species, light and water as
source factors) with Type III sums of squares (Quero et al.
M. micrantha -FI
M. micrantha -MI
M. micrantha -LI
C. odorata -MI
C. odorata -LI
16
12
8
FW
MW
LW
4
0
-4
(b) 20
C. odorata -FI
16
12
8
4
0
0
-4
-2 -1
123
73
Results
General effects of light and water
The reduction in the availability of light and water imposed
structural changes on the leaves of both species and
affected their physiological performance (Fig. 1, Table 2).
Most variables showed strong light effects and were species-dependent (Table 2). In comparison, the effects of dry
Table 2 Results of three-way ANOVAs for some structural and physiological traits, according to the treatment factors species (S), light (L) and
water (W)
Factor
R2
Interactions
Species
Light
Water
21.7***
42.2***
0.9***
S9L
S9W
L9W
S9L9W
3.4***
6.4***
1.4***
Structural traits
Area (cm2)
2
-1
SLA (cm g )
14.3***
6.3***
82.2
74.2***
0.7***
1.2***
0.7***
0.7***
1.0***
92.3
24.7***
3.7***
28.3***
1.0
5.6***
5.5***
69.0
16.5***
1.5*
25.1***
1.8*
5.4***
5.1***
65.1
62.2***
0.0
3.8***
1.2*
2.4**
2.3**
72.3
5.0**
16.2***
1.0
4.0*
3.0
53.1
5.5*
1.1
28.6***
54.6***
0.6
LMF (g g-1)
0.1
SMF (g g-1)
9.7***
0.2
4.6***
19.3***
Physiological traits
U (mol CO2 mol-1 photons)
0.4
h ()
14.2***
14.0***
1.8***
25.3***
3.5
79.2***
10.1***
0.3
4.7*
16.6***
0.0
3.3*
0.0
9.2***
5.2*
4.5
5.3
50.5
1.7
0.9
8.4***
80.7
4.9*
3.2
5.7*
52.6
2.9***
0.6
2.7**
0.3
3.0***
6.6*
89.5
55.4
1.8***
89.7***
0.0
0.6***
0.5**
0.2
1.8***
94.6
0.2
56.0***
2.8**
7.4***
1.2
2.4
7.1***
80.4
Ci/Ca
2.0**
56.3***
3.2***
12.1***
3.5***
2.7**
4.0**
83.7
5.1***
76.7***
2.5***
2.9***
1.2***
3.5**
3.0***
94.8
1.6***
78.8***
1.2***
4.9***
0.4*
1.9***
7.5***
96.2
The proportion of the explained variance (SS9/SStotal) and the level of significance for each factor and the interactions are indicated. R2 is the
proportion of total variance absorbed by the model. Area Single leaf area, SLA specific leaf area, LMF leaf mass fraction, SMF stem mass
fraction, RGR relative growth rate, Nmass nitrogen concentration, U apparent quantum yield, h bending degree or curvature, LCP light
compensation point, LSP light saturation point, Rd dark respiration rate, Pmax maximum light saturated photosynthetic rate, WUE water use
efficiency, Ci/Ca ratio of intercellular to external CO2 concentration, gs stomatal conductance, PNUE photosynthetic nitrogen-use efficiency
* P \ 0.05; ** P \ 0.01; *** P \ 0.001
123
74
20
M . micrantha
(a)
Aa
BaBa
15
Ab Ab
Ab
10
C . odorata
(b)
Ba
Ba
Aa
FW
MW
LW
Ab
Ab Ab
Ac
AcAc
Ac
Ac Ac
Area (cm2)
25
210
180
150
120
90
60
30
0
Ab Ac
(c)
Ba
Aa Aa
1200
Ac(d)
Ab
Aa
Aa
Aa
Aab
Ab
Aa Ab
Ab
Ab
Ba Aa
(mmol H2O)-1)
90
80
70
60
50
40
30
20
10
0
1000
800
600
M . micrantha
(a)
C . odorata
FW
MW
LW
Aa
AbAa
Ab
Ba
Ca
(b)
Ba
Aa
Ca
Ba
Bb
Ab t
Ab
Ac
Bb
AcAbAc
(c)
Ac Ac
Bb
Aa
Aa Ab
Aa
(d)
Ac
Bb
Ac
Bc Bc
Ab
Bb Cb
Aa
Aa Aa
400
200
60
50
40
(e)Aa
0.7
(f)
Ab Ab
0.6
Aa
Aa
Aa Aa Aa Ab
Bb
Ba
Ab Ac
Ac
AbAcAb
LMF(g g-1)
Aa
(g)
Aab
AbBbAb
Ba
Ab Aab
Aa Ba
Aa
Aa Aa
AbAaAa
20
10
Aa (i)
Ab
Ab
Ab
(f)
Aa Aa
Aa
Bab Bb Aa Bc
Aa
Aa
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
0.05
BaBa
Ab
Ab Aa
AaBa
Ca
Aa Aa Ab
Ab
Ab
Ac
200
AcAc Ab
100
0
FI
(g)Aa Aa Aa Aa Aa
Aa
(h)
AbAb
Ab
Aa
Aa
Aa Aa AaAaAab Aa
Bb
(j)
400
MI
LI
FI
MI
LI
RGR (g.g-1d-1)
300
0.4
0.3
0.2
Aa Aa
Aa
(h)
AB a
Aa Ba
30
500
0.5
(e)
Aa Aa
Aa
0.1
0.0
SMF(g g-1)
0
0.35
0.30
0.25
0.20
0.15
0.10
0.05
0.00
0.04
0.03
(i)
0.02
Ab Ab
Bc
Ab
0.01
(j)
AaAa Aa
Aa
Aa
Aa
Aa
AbAa
Ab
Ba
Ac
Ac Ac
FI
MI
LI
FI
MI
LI
123
75
(Fig. 3c,d). LMF and SMF were less affected by the varied
light and water interactions, particularly in C. odorata
(Fig. 3eh). Under the same light level, most structural
variables for both species did not vary significantly with
different water treatments (Fig. 3). Under LI, both species
had increased RGR with increasingly dry conditions, a
trend that was particularly significant for M. micrantha
(Fig. 3i, j).
Discussion
Response to variation of light and water
The response of both species to light (Responselight) was
generally higher than their response to water (Responsewater) (Fig. 4). Responselight in M. micrantha had a mean
value of 0.571 for the structural traits variables, which was
higher than that in C. odorata (Responselight = 0.533).
Conversely, the mean Responselight for the physiological
traits was 0.739 in M. micrantha, lower than that in
C. odorata (Responselight = 0.792). In particular, RGR of
C. odorata was strongly affected by light (Responselight = 0.989), suggesting that high light favored growth of
C. odorata. Both species showed the lowest response to
light and water in Nmass (Fig. 4).
Relationships between structural
and functional variables
The relationships between leaf structural and physiological
variables were similar for the two invasive weeds, with
integrated results shown in Table 3 and some separated
results in Fig. 5. The results indicated that most of the
correlations between structural and physiological variables
in both species did not change significantly with light
variance (data not shown). In most cases, the correlation of
structural traits with leaf physiological performance differed depending on water treatment (Table 3, Fig. 5).
Photosynthetic rates on a mass basis (Amass) were significantly correlated with RGR under FW and MW for M.
micrantha, and under all water treatments for C. odorata
C.odorata
M. micrantha
1.0
Structural
Physiological
0.8
1.0
RGR
Rd
LCP
0.8
0.6
LSP
Ci/Ca
WUE
Rd
0.4
PNUE
SLA
Pmax
SMF LMF
0.2
Response water
gs
0.0
RGR
Area
0.6
LCP
gs
Ci/Ca
SLA
0.4
WUE
LMF
PNUE
Pmax
SMF
Nmass
0.2
Nmass
LSP
Area
Response water
0.0
0.0
0.2
0.4
0.6
Response light
0.8
1.0
0.0
0.2
0.4
0.6
0.8
1.0
Response light
123
76
Table 3 Spearmans correlation coefficients for some log structural and physiological traits at three different water treatments on integrated data
from both weeds
Pmax
gs
PNUE
Nmass
SLA
RGR
LMF
SMF
WUE
LSP
FW
0.94**
0.89*
0.83*
-0.49
-0.89*
0.89*
-0.49
0.38
-0.37
MW
0.60
0.60
0.54
-0.14
-0.60
0.54
-0.54
0.37
-0.83*
LW
0.94**
0.60
0.77
-0.54
-1.00**
0.94**
-0.52
0.43
0.89*
WUE
FW
-0.31
-0.49
-0.49
0.14
0.20
-0.14
0.83*
-0.83*
MW
LW
-0.43
-0.49
-0.68
-0.83*
-0.49
-0.37
0.26
0.60
0.43
0.60
-0.37
-0.54
0.60
0.77
-0.37
-0.71
FW
0.2
0.41
0.26
-0.64
-0.06
0.03
-0.99**
MW
0.09
0.54
0.37
-0.77
-0.09
-0.03
-0.94**
LW
0.54
0.89*
0.66
-0.89*
-0.43
0.49
-0.93**
-0.14
SMF
LMF
FW
-0.31
-0.54
-0.37
0.66
0.14
MW
-0.26
-0.60
-0.43
0.83*
0.26
-0.09
LW
-0.67
-0.90*
-0.70
0.99**
0.52
-0.46
RGR
FW
0.83*
0.83*
0.66
-0.14
-0.89*
MW
0.94**
0.71
0.89*
-0.09
-0.94**
LW
0.89*
0.54
0.83*
-0.49
-0.94**
SLA
FW
MW
-0.94**
-1.00**
-0.66
-0.66
-0.83*
-0.83*
0.14
0.31
LW
-0.94**
-0.60
-0.77
0.54
Nmass
FW
-0.31
-0.37
-0.20
MW
-0.31
-0.43
-0.37
LW
-0.71
-0.83*
-0.77
PNUE
FW
0.94**
0.71
MW
0.83*
0.94**
LW
0.89*
0.54
gs
FW
0.77
MW
0.66
LW
0.66
123
77
M . micrantha
C . odorata
1200
(a)
1000
(b)
800
600
400
200
FW: r=0.79**
FW: r=0.50*
MW: r=0.84**
MW: r=0.77**
LW: r=0.40ns
LW: r=0.86**
0
0
0.02
0.04
0.06
0.02
0.04
0.06
RGR (g g-1d-1)
RGR (g g-1d-1)
0.05
(c)
0.04
(d)
FW: r= - 0.83**
FW: r= - 0.87**
MW: r= - 0.87**
MW: r= - 0.88**
LW: r= - 0.83**
LW: r= - 0.63**
0.03
0.02
0.01
0
400
800
1200
1600
400
800
1200
1600
(f)
(e)
400
(mol N)-1s-1)
500
300
200
100
FW: r=1.00**
FW: r=0.95**
MW: r=0.99**
MW: r=0.97**
LW: r=1.00**
LW: r=1.00**
0
0
500
1000
1500
2000
500
1000
1500
2000
Conclusions
The results presented here indicate that both species favor
high light conditions, while differing in their soil water
preferences: M. micrantha was particularly efficient at full
123
78
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