Journal of Horticultural Research 2016, vol.

24(1): 79-91
DOI: 10.1515/johr-2016-0010
_______________________________________________________________________________________________________

COMPETITIVE ABILITY OF CAPSICUM ANNUUM L. RELATIVE

TO THE WEED AMARANTHUS LIVIDUS L.
Abdessatar OMEZINE1*, Jaime A. TEIXEIRA DA SILVA2**
1

Institution of Agricultural Research and Higher Education, Tunis, Tunisia

2
Retired, Miki-cho, Japan
Received: September 2015; Accepted: April 2016

ABSTRACT
Amaranthus lividus is the most frequently reported troublesome weed in the production of Capsicum
annuum in some regions because it is an aggressive invader, difficult to control, and reduces yield significantly. The effects of A. lividus on the growth of C. annuum Baklouti were evaluated under greenhouse
conditions. Three experiments were conducted to determine the effect of A. lividus on the biomass accumulation of C. annuum seedlings for 40 days. In an additive experiment, to one C. annuum seedling and 1, 2, 3
or 4 A. lividus seedlings were grown per pot. The second experiment was established to assess C. annuum
intracompetition using from one to six plants per pot as the density. In a replacement experiment, C. annuum
and A. lividus seedlings were transplanted into pots with different proportions of both plants (1/5, 2/4, 3/3,
4/2, 5/1). Competition by A. lividus reduced C. annuum plant dry weight by as much as 93%. However,
C. annuum had little effect on A. lividus, reducing dry weight by 31.3% at a 5:1 ratio of C. annuum: A. lividus. Relative yield analysis between C. annuum and A. lividus demonstrated the competitive advantage of
C. annuum over A. lividus. The relative crowding coefficient of both plants changed significantly in the
presence of the other plant, at any ratio. The aggressivity of C. annuum was higher at its lower proportion
and C. annuum was more aggressive towards itself than towards A. lividus when its density increased.
Key words: aggressivity; allelopathy; competition; relative crowding coefficient; relative yield; weeds
INTRODUCTION
In Tunisia, Capsicum annuum L. (Solanaceae),
or peppers in general, is an important commercial
crop grown on a wide range of soils at altitudes varying from sea level to 2000 m, both for internal consumption and for export. C. annuum is ranked fifth
(in terms of acreage) after tomato, watermelon, potato and onion (Boughalleb & El Mahjoub 2005). In
Africa, Tunisia is the third largest producer of
C. annuum after Nigeria and Egypt, and the third
largest exporter (in terms of tonnage) after Morocco
and South Africa (FAO 2013). In 2008 and 2010,
Tunisia was ranked 12 and 11, in terms of global
production with a world rank of 45 and 53, respectively in export terms (FAO 2013).
The yield of C. annum in Tunisia averages
12.5 t/ha, which is relatively low compared to yield

*Corresponding authors:
e-mail: *abdessatar.omezine@yahoo.fr; **jaimetex@yahoo.com

observed in other Mediterranean countries such as

Spain (35 tha-1), Italy (28 tha-1), Greece (23 tha-1),
and Morocco (14 tha-1) (Boughalleb & El Mahjoub
2005). These low yields are due to difficulties in establishing efficient control measures. One of the
main problems affecting crop yield and quality is
weed interference (Hager et al. 2002; Boatright &
McKissick 2004). Severe weed infestations can reduce yield by at least 50-87% and some weeds,
among them Amaranthus lividus, may act as alternate hosts of insects, pests and diseases (Boatright
& McKissick 2004). Effective weed management is
thus one of the many critical components of successful C. annuum production.
The genus Amaranthus is represented by approximately 60 species (Holm et al. 1997;
Wiersema & Leon 2013), widely distributed in tropical, subtropical and temperate regions (Carvalho et

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al. 2006). A. lividus is among the most abundant

weeds occurring in and affecting vegetable crops,
lawns, pastures, gardens and row crops (Murphy et
al. 2010) throughout the warm temperate and tropical
regions of New and Old Worlds (Wiersema & Len
2013). It is responsible for significant yield losses in
many crops, which varies with the density of A. lividus. For example, the yield of C. annuum was reduced by 67% when it competed with four A. lividus
plantsm-2 (Morales-Payan et al. 2002). In another
uncontrolled study, three A. lividus plantsm-2 interfered with C. annuum, reducing yield by 33% (Morales-Payan et al. 2003). Maximum yield loss was
65% when six A. lividus plants were grown per m2
or 30% with a single A. lividus plantm-2 (MoralesPayan & Stall 2002). Those studies, however, failed
to assess the ratio of C. annuum: A. lividus (i.e., the
density of one species relative to the other), and thus
served as the basis for this study with the hypothesis
that the growth and successful survival of one species would change in the presence of the other.
In Tunisia, and even around the world, limited
information is available on chemical and biological
weed control, including that of specific weeds like
A. lividus (Murphy et al. 2010). However, the only
measure to control A. lividus and other weeds in vegetable crop stands, including those of C. annuum,
particularly in low-income countries, is cultural
methods, particularly periodic hoeing and hand pulling throughout the growing season. This process is
laborious, starts early in the season when the weeds
are small and the process is frequently repeated
throughout the season. Thus, to find a mechanism
that is able to limit particular plants growth using
plant density could theoretically reduce, or remove,
the need for chemical control, and serve as a practical,
low-cost technique for rural farmers globally. If our
first hypothesis held true (i.e., that C. annum could
negatively influence the growth of A. lividus), it then
becomes important to determine the level of interaction when the ratio of each was varied.
Except for the Morales-Payan-associated papers listed above, no systematic research has been
conducted thus far to explore the competitive behavior of C. annuum towards weeds. Against this
background, lack of detailed information about the
effect of A. lividus on the growth of C. annuum and

losses caused to growth, and the rapid spread of

A. lividus in vegetable production systems in Tunisia, North Africa and elsewhere around the world,
this study set out to address three key objectives,
conducted in three separate experiments: (1) to examine the effect of A. lividus density on C. annuum
growth during the seedling stage, which is the most
sensitive growth stage; (2) to obtain the appropriate
plant density of C. annuum to minimize intracompetition; (3) to test the competitive ability of C. annuum towards the weed A. lividus using a replacement method. The hope was that a suitable C. annuum : A. lividus ratio could be found to ensure
maximum weed control at no cost, since costs are
traditionally associated with chemical control or
weeding. To test these objectives, different competition indices were employed.
MATERIALS AND METHODS
To study inter- and intra-specific competition
between a crop C. annuum Baklouti and a weed
A. lividus, three pot experiments were performed under greenhouse conditions. They were conducted at
the Superior Institute of Agricultural Sciences (ISA)
of Chott-Mariem (Sousse, Tunisia) in the fall of 2010
and repeated in 2011. The experimental units were
plastic containers (8 cm in diameter and 10 cm deep)
filled with standard horticultural potting medium
(sand, manure, perlite; 1 : 1 : 1, v / v). Based on the
previous observations, this container size was chosen
to provide unrestricted C. annuum and A. lividus
growth for 40 days after planting. C. annuum seeds
were sown in standard horticultural potting medium
and transplanted at the true two-leaf stage into experimental containers. A. lividus seeds were collected
from local field stands in C. annuum fields near ISA.
A. lividus seeds were sown in experimental pots and
thinned after emergence to the numbers described in
Experiments 1 and 3. The pots were irrigated with
50 ml of potable water every two days from the start
to the end of the experiment. The salinity (mass of
dissolved salts) of the water was 1.2 g dm-3. After
40 days since capsicum transplantation, the whole
plant dry weight (DW) was determined as explained
next. The experimental design was completely randomized with five experimental units.

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In the additive experiment (Exp. 1), one C. annuum seedling was planted per pot, together with 1,
2, 3 or 4 A. lividus seedlings, which were spaced
2 cm apart of C. annuum seedling.
In the second experiment (Exp. 2), intracompetition between C. annuum seedlings was studied. In
one pot 1, 2, 3, 4, 5 or 6 plants were grown.
In the replacement experiment (Exp. 3), there
were six plants in one pot grown as a mixture of
C. annuum and A. lividus seedlings in different proportions. The following combinations were evaluated:
(1) 100% C. annuum (6 plants/pot, as monoculture);
(2) 83.3% C. annuum (5 plants/pot) + 12.7% A. lividus (1 plant/pot); (3) 66.6% C. annuum (4 plants/pot)
+ 33.3% A. lividus (2 plants/pot); (4) 50% C. annuum
(3 plants/pot) + 50% A. lividus (3 plants/pot); (5)
33.3% C. annuum (2 plants/pot) + 66.6% A. lividus
(4 plants/pot); (6) 16.6% C. annuum (5 plants/pot)
+ 83.3% A. lividus (5 plants/pot); (7) 0% C. annuum (i.e.,
an Amaranthus monoculture). Forty days after transplanting C. annuum, total dry weight (DW) of C. annuum and A. lividus were determined, as explained next.
At 40 DAP, C. annuum and A. lividus biomass
was separated. Roots of each plant were washed gently and thoroughly to remove soil particles so that the
root tissues remained intact. DW was determined by
drying the whole plant in an oven for 48 h at 80 C.
The relative performance of each species in the
A. lividus / C. annuum combination was calculated.
The relative yield (RY) of both species was analyzed
graphically as described by de Wit (1960) and Harper (1977). The RY of A. lividus (RYa), the RY of
C. annuum (RYc) and the total relative yield (RYT)
of both species were calculated separately according
to the following equations (Harper, 1977):
RYa = yield of A. lividus in the mixture / yield of
A. lividus in monoculture;
RYc = yield of C. annum in the mixture / yield of
C. annuum in monoculture;
RYT = RYc + RYa.
A value of RYT = 1 indicates that C. annuum
and A. lividus are demanding the same limiting resources, RYT > 1 indicates that the two species
make different demands on resources, so competition is avoided and RYT < 1 indicates that there is
mutual antagonism between C. annuum and A. lividus (de Wit & Goudriaan 1978).

The relative crowding coefficient (RCC) was

used to determine the competitive ability of a plant
to obtain limited resources when grown in a community setting compared to its ability to utilize those resources when grown in a monoculture (Aminpanah
2013). RCC was calculated for both species using the
formula of Hoffman and Buhler (2002):
RCC = [(DWc 1 5 / DWa 5 1) + (DWc 2 4 /
DWa 4 2) + (DWc 3 3 / DWa 3 3) +
(DWc 4 2 / DWa 2 4) + (DWc 5 1 /
DWa 1 5) + (DWc 6 0 / DWa 0 6)],
where DWc n n is the DW of C. annuum at a ratio
of n : n and DWa n n is the DW of A. lividus at
different ratios. Thus, for example, DWc 1 5 /
DWa 5 1 = the DW of one Capsicum plant and five
Amaranthus plants per pot divided by the DW of five
Amaranthus plants and one Capsicum plant per pot.
According to this definition, an RCC value > 1
signifies a competitive advantage for C. annuum
compared to A. lividus and the larger the RCC value,
the greater the competitiveness with C. annuum. In
contrast, an RCC value of < 1 indicates that A. lividus is more competitive than C. annuum. An RCC
value = 1 indicates that there is no competition or
competitive advantage or disadvantage between
both species. An increase in the RCC value for
a species as the proportion in the plant mixture increases indicates that the relative competitiveness of
that species has increased (Morales-Payan et al.
1999; Williams & McCarthy 2001; Zarochentseva
2012).
The third index assessed was aggressivity,
which is often used to determine the competitive relationship between C. annuum and A. lividus in a mixed
crop. The aggressivity of C. annuum (Ac) was calculated as follows (McGilchrist & Trenbath 1971):
Ac = (DW Capsicummix / DW Capsicummono)
(DW Amaranthusmix / DW Amaranthusmono),
where mix = mixture of both crops; mono = monoculture. If Ac = 0, then both crops are equally competitive; if Ac is positive, then C. annuum is dominant, and if Ac is negative, then C. annuum is weak.
Mean data sets of five replicates per treatment in
completely randomized design and repeated twice
were subjected to one-way analysis of variance
(ANOVA), and means were separated using
Fishers protected LSD at P = 0.05.

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RESULTS AND DISCUSSION

Additive experiment (Exp. 1)
Many factors interact to determine the outcome of competition between a weed and a crop.
Among these factors, weed population density is
a major factor. It is well known that crop density is
important in limiting the competitive effect of
weeds. Many other factors such as soil type and climate are beyond the control of farmers but crop density can be more easily controlled. This study aimed
to take advantage of this basic biological fact to try
and control a weed (A. lividus) in pepper production
systems using pot experiments. Seedlings in the
most sensitive growth stage of the plant were used.
A. lividus density had a significant effect on the
DW of C. annuum. DW accumulation of C. annuum
decreased as A. lividus density increased (Fig. 1). At
all densities, either one or four A. lividus plants negatively interfered with and reduced C. annuum DW.
This significant reduction of C. annuum DW
accumulation at all densities of A. lividus suggests
that the growth of C. annuum plants was retarded by
the concomitant growth of A. lividus. C. annuum
plants accumulated maximum DW (20.01 g/pot)
when grown without A. lividus while lowest DW
was obtained when one C. annuum plant was grown
with four A. lividus plants (0.46 g/pot). DW was reduced by about 80.8% due to competition from one
A. lividus plant and by 97.7% when A. lividus was
planted at a density of four plants. This relation implies that one A. lividus plant at lowest density had
a greater effect on C. annuum growth than one
A. lividus plant at a higher density. What is surprising is how so few A. lividus plants/pot were able to
reduce the growth of C. annuum.
It is likely that not only competition to nutrients but also other types of interactions, including
allelopathy, are involved. The study of Berry et al.
(2006) showed that the density of A. lividus, 1 to
2 plants/m2 , caused a 10% yield reduction of cucumber. Also, Morales-Payan and Stall (2002) reported that C. annuum yield was reduced by 67%
when competing with 6 A. lividus per square meter
of field throughout an entire season. These effects
of A. lividus on the DW of C. annuum reflect the

competitive ability of A. lividus (Procpio et al.

2004) and also its aggressiveness (Silva et al. 2009).
However, the accumulation of A. lividus DW
was modified by its density and was significantly
influenced by the presence of C. annuum. The highest accumulation of A. lividus DW was observed
with four A. lividus plants associated with one
C. annuum plant (52.33 g/pot) while the lowest DW
accumulation (35.77 g/pot) occurred when one
A. lividus plant grew in the presence of one C. annuum plant.
The DW accumulation of individual C. annuum plants decreased as A. lividus density increased but also, the DW accumulation of individual A. lividus plant decreased when its density increased (Fig. 2). Moreover, the total mean
DW / plant (C. annuum + A. lividus) decreased with
increase of A. lividus density (Fig. 2). A similar relation was found by Ronchi and Silva (2006) in experiment with young coffee plants growing with
four weeds.
Intraspecific competition experiment (Exp. 2)
In the intraspecific experiment, an increase in
C. annuum density increased the DW accumulation/pot up to five plants/pot, but decreased rapidly
at six plants/pot while the DW accumulation per individual plant was not influenced (Fig. 3), which is
a typical reaction concerning yield of total biomass.
Agarwal et al. (2007) noted that the yield of C. annuum fruit increased with its population growth up
to 120,000 plants/ha resulting in the highest marketable yield but not influencing fruit mass; however,
exceeding this population in the field, fruit yield decreased significantly. Jolliffe and Gaye (1995) reported that leaf area, leaf DW and shoot DW of
C. annuum decreased significantly as plant population reached 11.1 plants/m2.
In our study of intraspecific competition, the
observed increase in DW was not proportional to
density. At a high density of C. annuum (6 plants/pot),
the total DW of C. annuum was 164.4 g/pot whereas
at a low density (1 plant/pot), total DW of C. annuum
was 36.8 g/pot. In other words, a dry weight of one
C. annuum plant was 27.4 g when the density was
6 plants/pot and 36.8 g when the density was
1 plant/pot. Jolliffe and Gaye (1995) showed that

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Dry weight acumulation of each

species per pot (g)

a high population (11.1 plants/m2) of C. annuum decreased biomass accumulation per unit land area.
However, Harper (1977) found that the total DW of
Amaranthus retroflexus / pot was relatively constant but DW/plant decreased as A. retroflexus density increased.
Density affects plant architecture, alters growth
and developmental patterns and influences carbohydrate production and partitioning. Plant density may
influence the development and growth of vegetable
crops, including onion (Asaduzzaman et al. 2015).
The plasticity of plants is the capacity for marked
variation in an individual plant phenotype as a result
of environmental influences on that genotype during
development in which an individual plant can grow
larger or smaller depending on the resources available to it in its habitat (Price et al. 2003; Dekker 2011).
High densities decreased the absolute growth
rates of C. annuum but promoted shoot biomass accumulation per unit land area (Jolliffe & Gaye 1995).
Spehia et al. (2014) noted that plant height of C. annuum changed significantly in response to spacing:
closer spacing resulted in maximum plant height, but
wider spacing resulted in the highest number of fruits.
Quinto and Barraza (2009) found that phenotypic plasticity of C. annuum can be affected by
treatments, since a significant effect of plant density
was detected on stem diameter, leaf area and leaf
number, but plant height was less plastic. In their
study, stem DW, leaf DW, fruit DW and total DW
per plant decreased as plant density increased. This
phenotypically plastic ability of C. annuum based on
morphological traits (Lahbib et al. 2012) allows them
60

Cap/pot
Am/pot
Total/pot

50
40
30

to rapidly adapt to changing environments and enables them to continue to survive and reproduce across
variable environments (Ttard-Jones et al. 2011).
Replacement experiment (Exp. 3)
1. Dry weight accumulation
The DW accumulation of C. annuum (Cap/pot)
increased with the number of pepper seedlings/pot
(and proportion to A. lividus) A similar trend concerning number of plants and proportion to C. annuum was shown for A. lividus (Fig. 4). The highest
DW per pot was at highest proportion of pepper in
the mixture. The accumulation of DW by individual
C. annuum plants increased as its ratio in the mixture decreased. A similar trend was observed for
A. lividus in the mixture. The DW accumulation of
individual plants, either Capsicum or Amaranthus,
changed as their ratio in the mixture changed: from
1.43 to 3.58 g when the proportion of Capsicum
changed from 5 to 1 plants/pot and from 6.37 to
1.72 g when the proportion of Amaranthus changed
from 1 to 5 plants/pot.
Thus, as the density of C. annuum increased, it
inhibited the growth of its own population more than
it inhibited the growth of A. lividus population. The
same was true for A. lividus representing a classic case
of mutual antagonism. The reduction of DW of either
of C. annuum or A. lividus as its density increased
might be due to the production of allelochemicals by
C. annuum (Radhouane & Rhim 2014) and/or A. lividus (Jala & Wongsarasin 2012).
The DW calculated for plants of both species
did not depend on their mutual proportions (Fig. 5).

c
d

20
b

10

0
0

Density of Amaranthus lividus (number of plants per pot)

Fig. 1. Effect of density of Amaranthus lividus grown in the pots with one Capsicum annum plant on the dry weight (DW) of
C. annuum and A. lividus accumulated during 40 days, per pot. Data was separated by ANOVA, and significant differences
assessed by the F-test and LSD at P = 0.05, and indicated by different letters across densities. Bars represent means SE (n = 10)

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A. Omezine, J.A. Teixeira da Silva

Dry weight accumulation of

Capsicum and Amaranthus per
plant (g)

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40

Cap/plant
Ama/plant
Total/plant

35
30

b
a

25

20

c
d

15
10

0
0

1
2
3
Density of Amaranthus lividus (number of plants per pot)

Fig. 2. Effect of density of Amaranthus lividus grown in the pots with one Capsicum annuum plant on the total dry
weight (TDW) of C. annuum and A. lividus accumulated during 40 days, on a per plant basis. Data was separated by
ANOVA, and significant differences assessed by the F-test and LSD at P = 0.05, and indicated by different letters
across densities. Bars represent means SE (n = 10)
Dry weight accumulation of
Capsicum per plant and pot (g)

250

a
DW/plant
DW/pot

200

b
c

150
d
100
a

50

0
1

Density of Capsicum annuum (number of plants per pot)

Fig. 3. Dry weight (DW) accumulation of Capsicum annuum during 40 days on a per plant and on a per pot basis as
affected by plant density. Data was separated by ANOVA, and significant differences assessed by the F-test and LSD
at P = 0.05, and indicated by different letters across densities. Bars represent means SE (n = 10)
Dry weight accumulation of each
species in the mixture per pot (g)

16

12
10

Cap/pot
Ama/pot
Total dry/pot

14

bc

cd

f
b

f
6

e
d

e
f

4
2
0
c6a0

c5a1

c4a2
c3a3
c2a4
Proportion of each species in the mixture

c1a5

c0a6

Fig. 4. Dry weight (DW) accumulation in Capsicum annuum and Amaranthus lividus plants grown in the mixture,
expressed on a per-pot basis. c and Cap = Capsicum, Ama and a = Amaranthus. Data was separated by ANOVA, and
significant differences assessed by the F-test and LSD at P = 0.05, and indicated by different letters across proportions.
Bars represent means SE (n = 10)

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Competition between Capsicum annuum and Amaranthus lividus

Dry weight accumulation of each

species in the mixture in gram per pot

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8
Cap/plant
Ama/plant
DW/plant

6
5
a

3
e

f
a

1
0
c6a0

c5a1

c4a2
c3a3
c2a4
Proportion of each specisin the mixture

c1a5

c0a6

Fig. 5. Dry weight (DW) accumulation in Capsicum annuum and Amaranthus lividus plants grown in the mixture,
expressed on a per-plant basis. c and Cap = Capsicum, Ama and a = Amaranthus. Data was separated by ANOVA,
and significant differences assessed by the F-test and LSD at P = 0.05, and indicated by different letters across proportions. Bars represent means SE (n = 10)

2.

Relative yields
The relative yields of C. annuum (RYc / pot
and RYc / plant) and A. lividus (RYa / pot and
RYa / plant) and the relative total yield (RYT / pot
and RYT / plant) obtained from the replacement series are shown in Fig. 6 and 7. These parameters
provide support for the competitive interaction of
each species. These yield parameters were significantly affected by the mixture ratios.
The relative yield (RYc/pot) of C. annuum increased as its ratio decreased from 0.46 at 5 C. annum
plants to 0.85 at 1 C. annuum plant (Fig. 6). RYa/pot
decreased from 0.79 to 0.37 when the proportion of
A. lividus plants in the mixture increased from 1 to
5 plants/pot. The RY of each species was < 1, indicating that there was mutual antagonism between
both species. In this case, it is difficult to determine
if the species were antagonistic or whether both were
competitively using the same resources since RYT
obscures the behavior of each species. Future experiments that examine nutrient levels in the soil and
plants at each ratio could elucidate this mechanism.
The RYT (RYc + RYa) of the mixtures did not
change significantly with each mixture combination
(Fig. 6). This value greater than unit indicates that the
two species (C. annuum and A. lividus) used available resources more efficiently than expected based on
their respective yields when species were considered
individually. Values > 1 indicate that while two species compete for different resources, there is also

probably some degree of resource complementarity

between them (Fetene 2013). When two competing
species share the same resource by occupying different areas or habitats, then spatial resource partitioning occurs (Yachi & Loreau 2007). The coexistence
of plant species may result from niche partitioning,
or from differences in resource requirements among
species while in the process of complementarity,
a more diverse plant community can use resources
more completely, and thus be more productive
(Fridley 2001). Differences in plant species richness
can affect ecosystem processes through partitioning
of resources, whereby plants in more diverse communities may increase total resource capture, and
thus increase net primary production.
Such complementarity of resource use may occur in space or time, or depend on the type of resource
(Ewel 1986). Species that are deeply rooted have
more access to water and nutrients, which are not
available to more shallowly rooted species (Johnson
et al. 2000). The number of C. annuum roots decreased with increasing depth and distance from the
stem (Gough 2001). From another perspective, C. annuum explored upper layers of the soil while A. lividus explored deeper layers (personal observation).
There was a competitive advantage of C. annuum over A. lividus when the ratio of C. annuum in
relation to A. lividus was either 5/1 or 4/2, resulting
in higher C. annuum RY per plant (0.79 and 0.36)
and decreased A. lividus RY (0. 09 and 0.13) (Fig. 7).

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Relative yield of each species

(g dry weight per pot)

This suggests that C. annuum growth can reduce the

growth of A. lividus when its proportion is 5/1 or 4/2
where the values are 0.85 and 0.35. For both species,
RY is < 1, which denotes that for both species interspecific competition was greater than intraspecific
competition. In such a case, the biomass of either
species is reduced in the presence of the other species (Williams & McCarthy 2001).
The relative total yield (g TDW/pot) was constant and greater than 1. However, the RY for both
plants was not constant and was < 1 (Fig. 6). RYc
followed the hypothetical line except at the ratio
c2a4, where it deviated from its hypothetical curve.
RYa was also < 1 but above the hypothetical line.
RYa deviated from the hypothetical line for at least
two proportions (c3a3, and c4a2). These results indicate, first, that the two species have different demands for resources, and second, that A. lividus was
more competitive than C. annuum when the density
of C. annuum was greater than three plants although
less than three C. annuum plants per pot made C. annuum more competitive than A. lividus. In such a case,
there was benefit to A. lividus and damage to C. annuum. Second, the competition between both plants
occurred for the same environmental resources,
which were used more efficiently by A. lividus.
A comparison between C. annuum and A. lividus shows that the relative yield of each plant and
relative total yield of C. annuum and A. lividus in
a competitive relationship were reduced as their proportion in the mixture changed, as verified by the
1,4

concavity of the RYc and RYa curves (Fig. 7). In addition, RYT was < 1.0, which indicates antagonist
competition (de Wit & Goudriaan 1978), which suggests interference caused by the allelopathic action.
Radhouane and Rhim (2014) reported that
C. annuum is toxic intra-specifically and inter-specifically against pear millet. Tsuchiya et al. (1994)
stated that the yields and quality of C. annuum in
Korea had been decreasing due to continuous cropping, suggesting that allelopathy may have accounted for this phenomenon since the water or organic solvent extracts of leaves, stems, roots and
soil in which plants had been cultivated inhibited
seed germination. Moreover, these authors reported
that the methanolic extracts of the stem and roots of
C. annuum strongly inhibited radicle growth, while
the methanolic extracts of leaves and roots and aqueous root extract inhibited hypocotyl growth of C. annuum seedlings. This allelopathic potential of different parts of the C. annuum plant may be applied to
weed management (Gonzalez et al. 1992, 1993).
Capsaicin, abundant in the Capsicum genus
(C. annuum, C. frutescens and C. chinense) (Al Othman et al. 2011), may act as an inhibiting allelochemical after being released into soil after the decomposition of senescent pepper tissues or by exudation from
their roots and its effectiveness increases as the dose
increases, although the effectiveness differs among
target plants (Kato-Noguchi & Tanaka 2003).
a

1,2
1

0,8
0,6

a
b

0,2

c
d

0,4

RYc/pot
RYa/pot
RYT/pot

0
c5a1

c4a2
c3a3
c2a4
Proportion of each species in the mixture

c1a5

Fig. 6. Relative yields (RY) in g, dry weight per pot of Capsicum annuum (RYc) and Amaranthus lividus (RYa) as well
as relative yield total (RYT) estimated from Capsicum and Amaranthus grown in competition in a replacement experiment. c = Capsicum, a = Amaranthus. Dashed lines refer to the relative hypothetical productivity when there is no
interference by one species with another. Data was separated by ANOVA, and significant differences assessed by the
F-test and LSD at P = 0.05, and indicated by different letters across proportions. Bars represent means SE (n = 10)

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RYc/plant
RYa/plant
RYT/plant

Relative yield of each species

(g dry weight per plant)

1,2
1

0,8
0,6

0,4
0,2

b
c

d
c

0
c5a1

c4a2

c3a3

c2a4

c1a5

Proportion of each species in the mixture

Fig. 7. Relative yields (RY) in g, dry weight per plant of Capsicum annuum (RYc) and Amaranthus lividus (RYa) as
well as relative yield total (RYT) estimated from Capsicum and Amaranthus grown in competition in a replacement
experiment. c = Capsicum, a = Amaranthus. Dashed lines refer to the relative hypothetical productivity when there is
no interference by one species with another. Data was separated by ANOVA, and significant differences assessed by
the F-test and LSD at P = 0.05, and indicated by different letters across proportions. Bars represent means SE (n = 10)
Table 1. Values of relative crowding coefficients (RCC) and aggressivity for the mixture of Capsicum annuum and
Amaranthus lividus as affected by plant density evaluated after 40 days after planting
Proportions of
Capsicum : Amaranthus
1/5 (16.6%)
2/4 (33.3%)
3/3 (50.0%)
4/2 (66.6%)
5/1 (83.3%)

RCC of
Capsicum annuum
5.10 1.21 a
2.65 0.95 b
2.09 0.81 c
1.24 0.62 e
1.38 0.35 d

RCC of
Amaranthus lividus
0.82 0.21 e
1.65 0.35 d
2.95 0.51 a
2.53 0.65 b
2.20 0.15 c

Means of aggressivity
1.88 0.11 a
0.38 0.09 b
0.70 0.12 c
2.00 0.15 d
5.71 0.21 e

Means SE in each column followed by same letters at superscripts are not significantly different at p = 0.05 based on LSD test.
Experiment was repeated twice.

3. Relative crowding coefficient

RCC (Table 1) was used to measure the competitiveness of A. lividus against C. annuum and the competitiveness of C. annuum against A. lividus. For C. annuum, RCC decreased from 5.10 to 1.24 as its proportion increased from 1Cap/5Al to 4Cap/2Al, respectively.
The RCC of the proportion 5Cap/1Al was slightly larger
than the proportion 4Cap/1Al. However, the RCC of
C. annuum at two proportions (1Cap/5Al and 2Cap/4Al)
was greater than that of A. lividus. The RCC of A. lividus increased from 0.82 to 2.95 as its proportion decreased from 1Cap/5Al to 3Cap/3Al, respectively but decreased from 2.95 to 2.20 as its proportion decreased
from 3Cap/3Al to 5Cap/1Al, respectively. However, the
RCC of A. lividus at three proportions (3Cap/3Al,
4Cap/2Al and 5Cap/1Al) were greater than that of C. annuum. Species with a higher RCC indicate that it is
more competitive, or the stronger competitor (Fischer

et al. 2001). When the proportions 1Cap/5Al, 4Cap/2Al,

C. annuum was a superior competitor than A. lividus.
However, at the proportions were 3Cap/3Al, 2Cap/4Al,
and 5Cap/1Al, A. lividus was a superior competitor than
C. annuum. A. lividus had lower RCC values than
C. annuum when its proportion was larger than
16.6%, indicating that A. lividus was less aggressive
towards C. annuum than vice versa (Table 1).
Aminpanah et al. (2012) showed that the RCC
values for aboveground DW, root DW, tiller number,
leaf area, and height were higher for two weed species (Echinochloa crus-galli (L.) P. Beauv. and Echinochloa oryzicola Vasinger) than cultivated Oryza
sativa L. Hashemi, indicating that both weed species were superior competitors to rice.
4. Aggressivity
The aggressivity of C. annuum (Table 1), which is
the difference between the relative dry yields of

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C. annuum and A. lividus, was significantly influenced by the ratio of the species combination. The
increase in the proportion of C. annuum from 1Cap/5Al
to 5Cap/1Al (16.6% to 83.3%) decreased the aggressivity of C. annuum from +1.88 to 5.71, respectively,
However, at the proportions of 1Cap/5Al and 2Cap/4Al
(16.6% and 33.3%), aggressivity was positive and at
the proportions of 3Cap/3Al, 4Cap/2Al and 5Cap/1Al (50%
to 83.3%), aggressivity was negative. The aggressivity values for C. annuum decreased with its proportion in a pot. It was dominant at 1 and 2 seedlings per
pot and not dominant at higher proportions. The aggressivity of C. annuum is modified by its own presence (autoaggressivity) as well as by the presence of
A. lividus. From these results, increasing proportion
of C. annuum in the mixture decreased its aggressivity and increased the aggressivity of A. lividus while
the higher rate of C. annuum reduced its aggressivity.
Wahla et al. (2009) showed that when the aggressivity value of one component is positive, then
that component is more competitive than the other,
and has a dominant effect (Bhatti et al. 2006). Our results support the findings of Sarkar and Chakraborty
(2000), Sarkar and Sanyal (2000) and Sarkar and
Kundu (2001), who reported the dominant effect of
sesame (Sesamum indicum L.) having a positive
aggressivity value when grown in association with
mungbean (Vigna radiata L.), mashbean (Vigna
mungo (L.) Hepper) and groundnut (Arachis hypogaea L.), respectively. The aggressivity of S. indicum + A. hypogaea at a 3 : 2 row ratio (Sarkar and
Chakraborty, 2000), of S. indicum + V. mungo at a 2 : 1
row ratio (Sarkar & Sanyal 2000), S. indicum + V. radiata at a 3 : 2 row ratio (Sarkar & Kundu 2001) had
a positive value of 0.37, 0.07 and 0.26, respectively.
From our results, the control of A. lividus is
highly recommended, in order to prevent its competition, probably for nutrients, water, and light, thus
reducing Capsicum growth. In this study, the effect
of A. lividus competition may have been overestimated due to pot size. Pot size probably contributed
to nutrient competition due to constraints in root
growth caused by a small soil volume. Markham
and Halwas (2011) found a significant reduction in
plant growth when soil volume was reduced. Moreover, plants grown with neighbors tend to be smaller

than plants grown alone (Laird & Aarssen 2005).

Moreover, taking into account that interference
among neighboring plants occurs after a specific
plant density has been reached (Aldrich 1987; Ronchi & Silva 2006), in addition to competition between plants, intra-specific competition among individuals of the same species almost certainly had
also occurred, mainly at higher densities. Under
field conditions, soil volume restriction to root
growth is probably much lower than that observed
in pots, so field trials would be required to test this
hypothesis and to see whether the same growth responses of both crops would result in the field.
Moreover, in Tunisia, A. lividus densities in Capsicum fields are usually much higher than those examined in this study. Hachem (2003) reported 4550 seedlings of A. lividus / 50 cm2 in the field where
C. annuum was cultivated. This density could lead to
a higher degree of competition reported in our study.
CONCLUSIONS
The harmful effects of A. lividus competition
on Capsicum plant growth varied greatly depending
on A. lividus density. These adverse effects of
A. lividus on Capsicum growth were brought about
probably through competition mainly for essential
nutrients and light. The increase in the density of
A. lividus significantly decreased the growth of
C. annuum seedlings. The results from the replacement series indicate that the competitive relation between C. annuum and A. lividus changed with the
proportion of each species in the mixture. A. lividus
was a superior competitor than C. annuum at c3a3,
c4a2 and c5a1. The same tendency was observed for
C. annuum, at c1a5 and c2a4, in which C. annuum
was a superior competitor than A. lividus. These results suggest that under competitive conditions,
C. annuum produces more than expected when it is
found at a lower proportion in the mixture thus intraspecific competition is more important than interspecific competition. The same can be said for
A. lividus, which suffered from competitive interference when it was more abundant in the mixture
(i.e., it suffered more from intraspecific competition
than from competition with C. annuum plants).
However, a single factor, such as proportion, is not

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enough to predict interference between two species.

Future studies should evaluate the effect of water,
light and nutrients on the co-growth of these species. Under the present experimental conditions,
when even one A. lividus plant was able to significantly decrease the dry weight of C. annuum seedlings, the control of A. lividus is highly recommended to prevent competition with C. annuum and
to increase the growth (and consequently productivity and yield) of C. annuum.
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