American Journal of Plant Sciences, 2013, 4, 1839-1845
http://dx.doi.org/10.4236/ajps.2013.49226 Published Online September 2013 (http://www.scirp.org/journal/ajps)
Do Higher Resource Capture Ability and Utilization
Efficiency Facilitate the Successful Invasion of Exotic Plant?
A Case Study of Alternanthera philoxeroides
Xuyan Geng, Shang Jiang, Bo Li, Xiaoyun Pan*
Coastal Ecosystems Research Station of Yangtze River Estuary, Ministry of Education Key Laboratory for Biodiversity Science &
Ecological Engineering, Institute of Biodiversity Science, Shanghai, China.
Email: *xypan@fudan.edu.cn
Received June 16th, 2013; revised July 20th, 2013; August 20th, 2013
Copyright © 2013 Xuyan Geng et al. This is an open access article distributed under the Creative Commons Attribution License,
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
ABSTRACT
We tested the hypothesis that introduced populations may have higher resource capture ability and utilization efficiency
than native ones of invasive plants. We compared ecophysiological traits including maximum photosynthetic rate (Pmax),
apparent quantum yield (Q), specific leaf area (SLA), photosynthetic energy use efficiency (PEUE), photosynthetic ni-
trogen use efficiency (PNUE), water use efficiency (WUE), mass-based and area-based leaf construction cost (CCmass
and CCarea), and mass-based and area-based leaf nitrogen concentration (Nmass and Narea) between native (Argentina) and
introduced (USA) populations of two varieties (North Apa and South Apo) of Alternanthera philoxeroides under com-
mon garden conditions in China. For Apo and Apa, Pmax, Q, Nmass and WUE were not significantly different between
native and introduced populations; introduced populations had significantly lower SLA and lower CCmas s but signifi-
cantly higher Narea and CCarea than native ones. For Apa, the introduced populations showed significantly lower PEUE
and lower PNUE while for Apo, PEUE and PNUE were not significantly different between native and introduced
populations. The results indicated that introduced populations of A. philoxeroides do not show higher resource capture
ability and resource utilization efficiency than their native ones in the common garden experiment, suggesting that these
traits may not necessarily contribute to successful invasion of invasive plants.
Keywords: Invasive Plant; Resource Use Efficiency; Construction Cost; Leaf Nitrogen
1. Introduction
One mechanism which has been frequently mentioned
and investigated empirically in the context of plant inva-
sions is that higher resource capture ability and utiliza-
tion efficiency may facilitate successful invasion [1-3].
Studies comparing leaf traits of invasive plants and na-
tive plants have shown that invasive species have higher
specific leaf area (SLA) [4,5], lower mass-based leaf
construction cost (CCma s s ) [4,6-8], higher photosynthetic
nitrogen use efficiency (PNUE) [3,4,9], higher photo-
synthetic energy use efficiency (PEUE) [8,10] and higher
water use efficiency (WUE) [9,11] than native species.
However, some studies have detected that there are no
significant differences in CCmass [12], PNUE [8] and
WUE [6,8] between invaders and natives. A recent study
also suggests that the populations in the introduced range
of an invasive species have evolved a higher PEUE and a
shorter payback time but not lower CCmass than those in
the native range [13].
Do exotic invasive plants have generally evolved higher
resource capture ability and utilization efficiency in their
introduced ranges? To answer this question, we com-
pared leaf-level physiological traits that are related to re-
source capture and utilization efficiency, e.g., maximum
photosynthetic rate (Pmax), apparent quantum yield, SLA,
PNUE, PEUE, WUE, area-based leaf construction cost
(CCarea) and CCmass, area-based leaf nitrogen contention
(Narea) and mass-based nitrogen contention (Nmass), be-
tween native (Argentina) and introduced (USA) popula-
tions of two varieties of Alternanthera philoxeroides (al-
ligator weed) under common garden conditions in Shang-
hai, China.
*Corresponding author.
Copyright © 2013 SciRes. AJPS
Do Higher Resource Capture Ability and Utilization Efficiency Facilitate the Successful Invasion
of Exotic Plant? A Case Study of Alternanthera philoxeroides
1840
2. Materials and Methods
2.1. Plant Species
Alternanthera philoxeroides (alligator weed), a clonal
herbaceous perennial native to South America, is distrib-
uted from Buenos Aires province to southern Brazil
(18˚S - 39˚S) [14]. It has been widely introduced to warm
temperate and subtropical humid areas in North America,
Australia and China [15]. At present, A. philoxeroides is
one of the most noxious weeds in wetlands and agro eco-
systems [16]. Many attributes have contributed to the in-
vasion success of A. philoxeroides in China, such as ra-
pid vegetative growth and reproduction [15], higher phe-
notypic plasticity than its native congener A. sessilis [16,
17] and adaptation to physical disturbances[18].
A. philoxeroides typically emerges from belowground
buds (on storage roots) in spring and then spreads vege-
tatively throughout a growing season, consequently form-
ing dense monospecific stands. It overwinters with stor-
age roots and rhizomes [18]. Although A. philoxeroides
may produce viable seeds, sexual reproduction contrib-
utes little to population regeneration due to extremely
low seed outputs and low germination rates [14,15]. In
contrast, vegetative propagation (with storage roots and
stems) is its primary regeneration strategy in the field
[14].
At least two varieties of A. philoxeroides are recog-
nized in Argentina [19]: the northern A. p. var. acutifo lia
(Apa) and the southern A. p. var. obtusifolia (Apo) [20].
The two varieties are distributed in different geographic
areas and habitat types (Figure 1(a); Flooding Pampa
grasslands vs. wetlands along the Middle Parana River),
and have evolved different leaf and stem morphologies
[19,20]. Both Apa and Apo have been introduced into
USA (Figure 1(b)) [21].
In 2003 and 2004 we sampled stem fragments of A.
philoxeroides from 16 populations: eight Argentina po-
pulations and eight USA populations (Figure 1). Both
Argentina and USA populations were sampled widely
across their distribution range. We collected 10 - 20 stems
for each population (more than 20 m apart from each
other to ensure that different clones/genets might be sam-
pled). All collected stems were cloned in a greenhouse
for more than 3 years to reduce environmental maternal
effects.
2.2. Experiment Design
The study was carried in August 2012 at the Experimen-
tal Field Unit of Fudan University, Shanghai, China,
which is an outdoor field previously used for other gar-
den experiments. The climate is humid subtropical, with
rainfall averaging 1160 mm per year and with mean
monthly temperatures ranging from 27.9˚C in July to
(a) Argentina
TU
CH FO
SA
BA
NC
GA
MS
AR
(b) USA
TA LA
FL
GULF OF MEXICO
Figure 1. Sample sites of two varieties, Apa (black circle)
and Apo (black triangle), of Alternanthera philoxeroides
from native (Argentina) and introduced (USA) range.
4.2˚C in January. In July 2011, we vegetative cloned 16
populations in experiment garden. Two weeks later we
selected four individuals (with 2 - 3 internodes and four
or six leaves) per populations, and planted them individu-
ally at a depth of 2 cm in round pots (diameter, 23 cm;
depth, 18 cm).We used nutrient soil (Beilei, Beilei Organic
Fertilizer Co., Ltd., Zhenjiang, China) with the content of
N, P, K 2% (dry weight basis), organic matter 35%
(dry weight basis), water 45%, and pH 5.5 - 6.5. All
pots were randomly arranged on a desk and re-random-
ized weekly to reduce position effects. Pots were watered
by hand every other day to keep the substrate moist.
We harvested all plant materials 8 weeks after planting.
Each plant was separated into leaves, stems, and roots.
We determined leaf area for each plant with a leaf area
meter (LI-3100A; LI-COR, Lincoln, NB, USA). All ma-
terials were oven dried at 55˚C for 72 h and then weighed
to the nearest 0.001 g. SLA (cm2·g–1) was calculated as
the ratio of leaf area to leaf dry mass. And then we finely
ground the dried leaves.
Mass-based carbon concentration (C) and Nmass of the
powdered leaves were determined with OEA analyzer
(Organic Elemental Analysis, FlashEA1112, Thermo Fin-
nigan, Italy). Ash concentration (Ash) was determined
after combusting leaf sample in a muffle furnace at
550˚C for 6 h. Ash alkalinity (AA) was determined aci-
dimetrically [22]. The mineral concentration (Min) of
Copyright © 2013 SciRes. AJPS
Do Higher Resource Capture Ability and Utilization Efficiency Facilitate the Successful Invasion
of Exotic Plant? A Case Study of Alternanthera philoxeroides
Copyright © 2013 SciRes. AJPS
1841
each sample was calculated according to [22] as follows:
MinAsh AA30 Nitrate, (1)
CCmass can be calculated [23,24] as follows:
C
CC1.041 5.077 1000 Min
1000 MinNorg
5.325 .
1000 1000
 





(2)
Narea and Nmass), with range (Argentina vs. USA) as a
fixed factor, population nested within range as a random
factor. The residuals for all tests were normally distrib-
uted and no transformations were necessary. All analyses
were carried out using SPSS 13.0 (SPSS, Chicago, USA).
3. Results
For both Apo and Apa, Pma x (Figure 2(a)) and apparent
quantum yield (Figure 2(b)) were not significantly dif-
ferent between native and introduced populations, and
introduced populations had significantly lower SLA
(Figure 2(c)).
Nitrate concentration were negligible, thus we as-
sumed that Norg = Nmass [25].
2.3. Determination of Physiological Traits For Apa the introduced populations showed signifi-
cantly lower PEUE and PNUE, while for Apo, PEUE and
PNUE were not significantly different between native
and introduced populations (Figures 3(a) and (b)). For
both Apa and Apo, WUE were not significantly different
between native and introduced populations (Fi gure 3(c )).
Net photosynthetic rate in relation to varying photosyn-
thetic photon flux density (light response curves) was
determined on the youngest fully-expanded leaves with a
Li-6400 Portable Photosynthesis System (LI-6400; LI-
COR, Lincoln, NB, USA). Measurements were made on
2 - 3 representative leaves on 4 individuals of each varie-
ties in a pair during continuously sunny days from 09:00
to 12:00 am in the field in August 2012. All leaves of the
Argentina and USA populations in a pair were of similar
age and position on the stems. PPFD decreased in a step-
wise fashion from 2000 to 0 µmol photon m–2·s–1 (at
2000, 1500, 1200, 1000, 800, 600, 400, 200, 150, 100, 70,
50, and 0). During the measurements, CO2 concentration,
temperature and relative humidity within the leaf cham-
ber were similar to those of ambient conditions. Each
leaf was acclimated for 10 - 20 min to 2000 µmol photon
m–2·s–1 of PPFD prior to the measurement.
For both Apa and Apo, Nmas s were not significantly
different between native and introduced populations (Fig-
ure 4(a)), but introduced populations had significantly
higher Narea than native ones (Figure 4). Introduced po-
pulations showed significantly lower CCmass (Figure
4(c)), but significantly higher CCarea (Figure 4(d)) than
native ones.
4. Discussion
4.1. Resource Capture Ability
Our results showed that introduced populations of A. phi-
loxeroides showed no significant difference in Pmax, ap-
parent quantum yield, and Nmass relative to the native po-
pulations, suggesting that invasive populations of A. phi-
loxeroides do not have advantage in resource capture
ability compared to their native populations. This result
was inconsistent with previous studies. Previous studies
have found that most invasive plants have higher Pmax
[6,9,11] compared to their co-occurring natives. Ref [8]
found that invasive species had significantly higher Pmax
(mass-based) than their non-invasive alien congeners. In
a study of comparing the functional traits between plants
from invasive and native populations of alien plant,
higher Pmax and higher SLA of invasive populations was
also found [13].
We fitted entire light response curves using the non-
rectangular hyperbola model according to [26] as shown
in Equation (3), where Photo is the leaf net photosynthe-
sis rate, PAR is the light intensity, Pmax is maximum
photosynthetic rate, Q is apparent quantum yield, R is
dark respiration rate and K is a constant.
Narea (g·m–2) = Nmass/SLA;
CCarea (g·glucose·m–2) = CCmass/SLA;
PEUE (µ·mol CO2 g glucose–1·s–1) = Pmax/CCarea [6];
PNUE (µ·mol CO2 g–1·s–1) = Pmax/Narea [27];
WUE (µ·molCO2 µmol–1 H2O) = Pmax/E (transpiration)
[9,28].
2.4. Statistical Analysis SLA can be envisaged as an indicator of thickness or
density; leaves with a higher SLA are typically thin and
have greater levels of herbivory [29]. Our results showed
that invasive plants of A. philoxeroides had significantly
lower SLA than their corresponding natives, indicating
Nested analysis of variance (ANOVA) was used to com-
pare if there had differences between plants from inva-
sive and native populations in parameters (Pmax, apparent
quantum yield, SLA, PNUE, PEUE,WUE, CCarea, CCmass,

2
max maxmax
PAR QPPAR QP4PAR QPK
Photo R.
2K
 
(3)
Do Higher Resource Capture Ability and Utilization Efficiency Facilitate the Successful Invasion
of Exotic Plant? A Case Study of Alternanthera philoxeroides
1842
(a)
N
ative Introduced
NS
NS
Apo Apa
NS NS
(b)
Apo Apa
(c)
Apo Apa
0
50
100
150
200
250
300
Specific leaf area (SLA)
(cm
2
·g
-1
)
0.00
0.02
0.04
0.06
0
Maximum photosynthetic rate
(μmol CO
2
m
-2
·s
-1
)
Apparent quantum yield
(μmol CO
2
per μmol photone)
5
10
15
20
25
Figure 2. Differences in maximum photosynthetic rate (Pmax
(a), apparent quantum yield (b) and speci fic leaf area (SLA)
(c) between native (Argentina) and introduced (USA) po-
pulations of two varieties (Apo and Apa) of Alternanthera
philoxeroides. Dates are means ± SE (NS, no significant dif-
ferences, *P < 0.05).
introduced populations of A. philoxeroides may have
evolved higher resistance to generalist herbivores than
native ones (Pan et al. unpublished results).
4.2. Resource Utilization Efficiency
Our results showed that invasive Apa had significantly
lower PEUE and PNUE than native Apa, and invasive
(a)
N
ative Introduced
NS
Apo Apa
PEUE
(μmol CO2 g·glucose-1·s-1)
0.4
0.3
0.2
0.1
0.0
(b)
NS
Apo Apa
0
3
6
9
12
15
PNUE
(μmol CO2 g-1·s-1)
(c)
NS
NS
Apo Apa
0
WUE
(μmol CO2 μmol-1 H2O)
1
2
3
4
5
6
Figure 3. Differences in photosynthetic energy use efficiency
(PEUE) (a), photosynthetic nitrogen use efficiency (PNUE)
(b) and water use efficiency (WUE) (c) between native (Ar-
gentina) and introduced (USA) populations of two varieties
(Apo and Apa) of Alternanthera philoxeroides. Dates are
means ± SE (NS, no significant differences, *P < 0.05 and
**P < 0.01).
Apo had no significant difference in PEUE and PNUE
compared to native Apo. These results indicate that in-
troduced populations of A. philoxeroides do not have
higher resource utilization efficiency than their native
ones in the common garden experiment. This contrasts
Copyright © 2013 SciRes. AJPS
Do Higher Resource Capture Ability and Utilization Efficiency Facilitate the Successful Invasion
of Exotic Plant? A Case Study of Alternanthera philoxeroides
1843
(b)
N
ative Introduced
NS
Apo Apa
N
area
(g·m
-3
)
0
NS
1
2
3
4
N
mass
(%)
0.0
0.5
1.0
1.5
2.0
(c)
(d)
Apo Apa
Apo Apa
Apo Apa
0.0
0.5
1.0
1.5
CC
area
(g·glucose·m
-2
)
CC
mass
(g·glucose·m
-1
)
0
10
20
30
40
50
60
70
80
Figure 4. Differences in mass-based nitrogen concentration
(Nmass) (a) area-based nitrogen concentration (Narea) (b)
mass-based construction cost (CCma ss ) (c) and area-based
construction cost (CCarea) (d) between native (Argentina)
and introduced (USA) populations of two varieties (Apo
and Apa) of Alternanthera philoxeroides. Dates are means ±
SE (NS, no significant differences, * P < 0.05, ** P < 0.01 and
***P < 0.001).
with previous studies. Invasive plants have often higher
PNUE [4,9,30], higher PEUE [8] or both higher PEUE
and PNUE [6,10,12] than natives species. Also in their
work comparing energy use strategy of an invasive spe-
cies from populations of its native ranges and introduced
ranges, Ref. [13] found that plants from invasive popula-
tions had a higher PEUE than native populations.
Our results showed that invasive populations of A. phi-
loxeroides had significantly lower CCmass and the same
levels of Nmass compared with native populations. How-
ever, due largely to significantly lower SLA, invasive
populations had significantly higher CCarea and Narea than
native populations (Figure 4). Nitrogen allocation to the
photosynthetic apparatus is suggested to be a major fac-
tor responsible for the interspecific variation in PNUE
[31]. Introduced Apa had the same leaf Nmass but signifi-
cantly lower PNUE than native ones, suggesting a de-
creased fraction of leaf nitrogen invested in the photo-
synthetic apparatus in introduced plants of Apa.
WUE did not differ between native and invasive po-
pulations of A. philoxeroides, which is accordant with
previous results that invasive plants are not at an advan-
tage over native species in WUE [6,8,9].
Our finding that introduced populations of A. philox-
eroides do not have higher resource capture ability and
resource utilization efficiency than their native ones in
the common garden experiment, suggesting that these
traits may not necessarily contribute to successful inva-
sion of invasive plants.
The importance of resource use efficiency will vary
across habitats and timescales [6]. Future research will be
needed to examine if there are differences in resource
capture ability and utilization efficiency between intro-
duced and native populations of invasive plants across
resource gradients and across their whole growth sea-
sons.
5. Acknowledgements
This study was financially supported by the National Na-
tural Science Foundation of China (31070369).
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of Exotic Plant? A Case Study of Alternanthera philoxeroides
1844
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Do Higher Resource Capture Ability and Utilization Efficiency Facilitate the Successful Invasion
of Exotic Plant? A Case Study of Alternanthera philoxeroides
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