How Do the Extracts From the Invasive Plant Acmella radicans Inhibit Germination and Growth of Brassica rapa (Field Mustard) and Chrysanthemum coronarium (Garland Daisy)? (2025)

International Journal of Agronomy

Volume 2025, Issue 1 8270456

Research Article

Open Access

This article is part of Special Issue:

Shicai Shen,

Shicai Shen

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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Gaofeng Xu,

Gaofeng Xu

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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Shaosong Yang,

Shaosong Yang

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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Guimei Jin,

Guimei Jin

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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David Roy Clements,

David Roy Clements

Department of Biology , Trinity Western University , Langley , British Columbia, Canada , twu.ca

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Kexin Yang,

Kexin Yang

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

School of Agriculture , Yunnan University , Kunming , China , ynu.edu.cn

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Xiaohan Wu,

Xiaohan Wu

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

School of Agriculture , Yunnan University , Kunming , China , ynu.edu.cn

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Zewen Fan,

Zewen Fan

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

School of Agriculture , Yunnan University , Kunming , China , ynu.edu.cn

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Fudou Zhang,

Corresponding Author

Fudou Zhang

  • [emailprotected]

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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Shicai Shen,

Shicai Shen

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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Gaofeng Xu,

Gaofeng Xu

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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Shaosong Yang,

Shaosong Yang

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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Guimei Jin,

Guimei Jin

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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David Roy Clements,

David Roy Clements

Department of Biology , Trinity Western University , Langley , British Columbia, Canada , twu.ca

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Kexin Yang,

Kexin Yang

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

School of Agriculture , Yunnan University , Kunming , China , ynu.edu.cn

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Xiaohan Wu,

Xiaohan Wu

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

School of Agriculture , Yunnan University , Kunming , China , ynu.edu.cn

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Zewen Fan,

Zewen Fan

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

School of Agriculture , Yunnan University , Kunming , China , ynu.edu.cn

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Fudou Zhang,

Corresponding Author

Fudou Zhang

  • [emailprotected]

Key Laboratory of Prevention and Control of Biological Invasions , Ministry of Agriculture and Rural Affairs of China , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

Yunnan Lancang-Mekong Agricultural Bio-Security International Science and Technology Cooperation Joint Research Center , Agricultural Environment and Resource Research Institute , Yunnan Academy of Agricultural Sciences , Kunming , China , yaas.org.cn

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First published: 08 May 2025

Academic Editor: Nidhi Chaudhary

Abstract

The allelopathic effects have not previously been studied for Acmella radicans (Jacquin) R. K. Jansen, a new invasive species recorded for Yunnan Province, China. In the current study, the allelopathic potential of root, stem, leaf, flower, and fruit head aqueous extracts from A. radicans on two important vegetable crops, Brassica rapa (field mustard) and Chrysanthemum coronarium (garland daisy), was explored in the laboratory. All four plant part aqueous extracts had some inhibitory effects on B. rapa and C. coronarium in terms of seed germination and seedling growth. Increasing concentrations of aqueous extracts of A. radicans reduced many indicators significantly, including germination rate, the seed germination index, root length, stem length, and biomass of B. rapa and C. coronarium. Generally, the most inhibited parameter was root length, followed by shoot length and biomass, with the lowest being seed germination. In terms of allelopathic response index, flowers and fruit heads yielded the strongest inhibition, followed by leaves and stems, with the lowest impact from root extracts. Root extracts had positive effects at 0.0125–0.025 g·mL−1. The inhibition rates for A. radicans extracts on B. rapa were generally higher than those of C. coronarium. It is concluded that A. radicans has allelopathic potential on vegetable crops with the overall inhibition rates ranked in the order flower and fruit head > leaf > stem > root. This was the first study to show that the allelopathic potential of A. radicans against crops may be strongly linked to its invasion and expansion.

1. Introduction

Alien plant species invasions pose significant threats to global biodiversity leading to various environmental and economic challenges [1, 2]. Their dominance in ecosystems arises from rapid growth, physiological and ecological adaptability, and often the use of allelopathic effects [36]. Such allelopathic impacts tend to increase the competitiveness of invasive plants [6, 7]. Plant allelopathy is the chemical interaction of one plant on other plants through the production of chemicals, which affects the seed germination, plant growth, and population establishment of the neighboring plants [8]. The majority (51.4%) of 524 invasive plant species could produce allelochemicals with the potential to negatively affect native plant performance [9], indicating that allelopathy could have a large impact on native species across the globe. Moreover, allelopathy is usually considered as one of the mechanisms by which such exotics may become successful invaders [69]. Hence, investigating the allelopathic potential of invasive plants is critical for understanding the potential competitive mechanisms employed by particular invasive plants against crops or native plant species.

The annual herbaceous plant Acmella radicans (Jacquin) R. K. Jansen originated in Central America [10]. Currently, the nonnative range of A. radicans includes Bangladesh, China, Cuba, Curaçao, India, Tanzania, and Thailand [1014]. In 2014, A. radicans was first documented as a naturalized species in China, specifically in Anhui Province [10]. This species thrives in moist weedy environments, including roadside ditches, riparian zones, and relatively damp agricultural fields like rice paddies. It is a relatively tall plant (up to 155 cm), with broad leaves and long sexual reproduction period (November through March), and produces a large number of seeds. Acmella radicans also has several medicinal applications, such as alleviating toothaches and treating infections [12, 15, 16].

Acmella radicans was identified as a potential concern in Yunnan Province, Southwestern China in 2017. By that time, it was already widely established in the vicinity of both Baoshan and Lincang, where it primarily invaded anthropogenically disturbed habitats like farm fields, tea plantations, orchards, roadsides, and ditches. The plant has rapidly become dominant in local habitats, adversely impacting plant species diversity and soil nutrients [17]. In an additional field survey, it has been found that A. radicans appeared to inhibit the growth of some vegetable species, especially two local major vegetable crops Brassica rapa (field mustard) and Chrysanthemum coronarium (garland daisy). Acmella radicans was widely distributed in fields of B. rapa and C. coronarium, with the crops exhibiting reduced seed germination rates and delayed germination compared to fields without A. radicans. Moreover, the yields of the two vegetable crops were also reduced when A. radicans was present. However, it was not known whether the impact of A. radicans on these and other species was related to allelopathic effects.

This study hypothesized that some of the observed impacts of A. radicans may be related to allelopathic effects. In order to explore the allelopathic potential of A. radicans against the two major vegetable crops B. rapa and C. coronarium, investigations were conducted to assess the allelopathic potential of aqueous extracts from root, stem, leaf, and flower and fruit heads of A. radicans to inhibit seed germination and growth of these vegetable crops.

2. Materials and Methods

2.1. Study Species

Acmella radicans plants were collected during the flowering and fruiting period in three regions (20°14′ N-25°12′ N, 99°16′ E−100°12′ E) in Yunnan Province on 24th December, 2022: Mengtong Township, Changning County, and Baoshan City. The plant samples were divided into four parts (root, stem, leaf, and flower and fruit head) after harvesting. The plant samples were dried at 40°C for a week and then pulverized with a grinder for preparation of extracts.

Seeds of the vegetable crops B. rapa and C. coronarium which exhibited germination rates over 95% in pre-experimental trials were acquired from a Changning County market in Yunnan Province for the bioassay experiment.

2.2. Extract Preparation and Bioassay

From January to March 2023, the allelopathic effects of root, stem, leaf, and flower and fruit heads of A. radicans on B. rapa and C. coronarium were tested under laboratory conditions. The root, stem, leaf, and flower and fruit head extracts for A. radicans were prepared using 50 g of fine powder that was dissolved at room temperature (20 ± 5°C) in 500 mL of distilled water (10% w/v). Before filtering takes place, the extracts were refrigerated for 48 h. Filtering was accomplished by pouring the extracts through cheese cloth (two layers) and filter paper (two layers). The initial aqueous extract concentration after filtering was 0.1 g·mL−1. Distilled water was used to dilute the initial concentration to form four concentrations (based on pre-experimental trials): 0.0125, 0.025, 0.05, and 0.1 g·mL−1, which were stored prior to use at 4°C.

Four extract concentrations (0.0125, 0.025, 0.05, and 0.1 g·mL−1) were tested, along with a control (distilled water) (CK), to investigate the potential allelopathic effects of A. radicans, with four replicates for each concentration. The bioassay procedures for assessing germination and seedling development of B. rapa and C. coronarium followed the methods outlined by Shen et al. [18]. After a 7-day period, root length, shoot height, and fresh biomass of the germinated vegetable seedlings were recorded.

2.3. Statistical Analysis

The experiments calculated germination rate (GR (%) = seeds germinated/total seeds × 100%), germination index (GI = Σ(GT/DT), where GT is the number of seed germinated on day T, and DT is the number of days from the beginning of the test) [19], and allelopathic response index (RI: when T ≥ C, RI = 1-C/T; when T < C, RI = T/C-1; C is the control value and T is the treatment value) [20] of B. rapa and C. coronarium subjected to aqueous extracts from A. radicans. Data were analyzed using two-way ANOVA to evaluate seed germination and growth parameters and one-way ANOVA to evaluate the allelopathic response index. Duncan’s multiple range tests were conducted to compare treatment differences (p < 0.05) for significant differences detected by ANOVA. The F statistic factors consisted of plant part and concentration level and their interaction (ns, , and ∗∗ indicate p > 0.05, p ≤ 0.05, and p ≤ 0.01, respectively).

3. Results

3.1. Germination Index and Germination Rate

Germination rate and germination index of B. rapa and C. coronarium varied significantly with plant part and concentration level of A. radicans extracts, with a significant interaction effect (p < 0.01) (Table 1 and Figure 1). Germination rate and germination index of both B. rapa and C. coronarium were significantly influenced by the aqueous extracts from all four plant parts of A. radicans tested (Table 1). These extracts exhibited powerful inhibitory effects on germination of the two test species (Table 1 and Figure 1). With increasing concentration, inhibition by the aqueous extracts was greatly increased, and germination of B. rapa was completely inhibited at concentrations of 0.05–0.1 g·mL−1 for either leaf or flower and fruit head aqueous extracts. For C. coronarium, the germination was completely inhibited by a 0.1 g·mL−1 leaf extract and a 0.05–0.1 g·mL−1 flower and fruit extract.

Table 1. Effects of aqueous extracts from four plant parts of Acmella radicans on seed germination of Brassica rapa and Chrysanthemum coronarium.
Plant part Concentration (g·mL−1) Brassica rapa Chrysanthemum coronarium
Germination index Germination rate (%) Germination index Germination rate (%)
Root 0.1 2.83 ± 0.17e 43.33 ± 2.72d 3.69 ± 0.21e 65.00 ± 4.30c
0.05 5.12 ± 0.28d 80.00 ± 2.72c 6.25 ± 0.37d 91.67 ± 4.30b
0.025 6.44 ± 0.22c 89.17 ± 4.19b 6.69 ± 0.19c 95.00 ± 4.30ab
0.0125 8.20 ± 0.33b 97.50 ± 3.19a 7.51 ± 0.17b 98.33 ± 3.33a
CK 8.83 ± 0.10a 98.33 ± 1.92a 8.23 ± 0.41a 98.33 ± 1.92a
  
Stem 0.1 2.75 ± 0.25e 42.50 ± 1.67d 3.33 ± 0.04e 60.00 ± 2.72c
0.05 4.77 ± 0.25d 71.67 ± 1.92c 5.55 ± 0.34d 86.67 ± 2.72b
0.025 6.19 ± 0.45c 85.00 ± 6.94b 6.62 ± 0.08c 96.67 ± 2.7a
0.0125 7.89 ± 0.21b 93.33 ± 2.72a 7.28 ± 0.59b 96.67 ± 4.71a
CK 8.83 ± 0.10a 98.33 ± 1.92a 8.23 ± 0.41a 98.33 ± 1.92a
  
Leaf 0.1 0.00 ± 0.00d 0.00 ± 0.00c 0.00 ± 0.00e 0.00 ± 0.00d
0.05 0.00 ± 0.00d 0.00 ± 0.00c 3.63 ± 0.22d 60.00 ± 4.71c
0.025 4.11 ± 0.33c 60.83 ± 3.19b 5.23 ± 0.14c 83.33 ± 2.72b
0.0125 7.60 ± 0.19b 97.50 ± 1.67a 6.91 ± 0.21b 96.67 ± 2.72a
CK 8.83 ± 0.10a 98.33 ± 1.92a 8.23 ± 0.41a 98.33 ± 1.92a
  
Flower and fruit head 0.1 0.00 ± 0.00d 0.00 ± 0.00c 0.00 ± 0.00d 0.00 ± 0.00d
0.05 0.00 ± 0.00d 0.00 ± 0.00c 0.00 ± 0.00d 0.00 ± 0.00d
0.025 3.83 ± 0.49c 60.00 ± 5.44b 2.69 ± 0.47c 43.33 ± 7.20c
0.0125 7.39 ± 0.52b 97.50 ± 1.67a 5.30 ± 0.26b 75.83 ± 4.19b
CK 8.83 ± 0.10a 98.33 ± 1.92a 8.23 ± 0.41a 98.33 ± 1.92a
  
F value Plant part 5.86∗∗ 669.78∗∗ 8.09 796.60∗∗
Concentration 32.26∗∗ 2332.31∗∗ 18.59 1019.12∗∗
Interaction 78.14∗∗ 170.74∗∗ 60.89 128.85∗∗
  • Note: Means with different letters are significantly different (p < 0.05).

3.2. Shoot Length and Root Length of Seedlings

Extracts of A. radicans affected the lengths of shoots and roots for both B. rapa and C. coronarium significantly (Figure 1 and Table 2). The aqueous extracts of A. radicans produced varied effects on the root and shoot lengths of the two vegetable crops (Table 2). Most extracts exhibited strong inhibitory effects on both species, with a few exceptions: root extracts of 0.0125 g·mL−1 for B. rapa as well as root extracts of 0.0125–0.025 g·mL−1 and stem extracts of 0.0125 g·mL−1 for C. coronarium showed stimulatory effects. Inhibitory rates of the aqueous extracts on root and shoot length of B. rapa and C. coronarium increased significantly with increasing concentration. Extracts from either flower and fruit head or leaf components of A. radicans generally had the strongest effect, with little or no growth of the bioassay species at the two highest concentrations, whereas the bioassay species still exhibited some growth at the higher concentrations of extract derived from roots or stems. The suppression rates of the aqueous extracts on B. rapa were generally greater than those on C. coronarium.

Table 2. Effects of four plant part aqueous extracts of Acmella radicans on the growth of Brassica rapa and Chrysanthemum coronarium.
Plant part Concentration (g·mL−1) Brassica rapa Chrysanthemum coronarium
Shoot length (cm) Root length (cm) Biomass (g) Shoot length (cm) Root length (cm) Biomass (g)
Root 0.1 0.19 ± 0.02e 0.49 ± 0.04e 0.029 ± 0.005d 0.23 ± 0.02e 0.44 ± 0.06e 0.012 ± 0.002c
0.05 0.26 ± 0.02d 0.59 ± 0.04d 0.033 ± 0.005d 0.42 ± 0.02d 0.55 ± 0.02d 0.013 ± 0.001c
0.025 0.45 ± 0.02c 0.92 ± 0.04c 0.091 ± 0.004c 0.58 ± 0.02c 0.79 ± 0.03c 0.017 ± 0.003b
0.0125 1.11 ± 0.11a 3.34 ± 0.03b 0.159 ± 0.007b 0.62 ± 0.02b 1.39 ± 0.05b 0.044 ± 0.001a
CK 0.93 ± 0.02b 3.49 ± 0.04a 0.191 ± 0.004a 0.55 ± 0.01a 1.78 ± 0.01a 0.041 ± 0.002a
  
Stem 0.1 0.17 ± 0.01e 0.43 ± 0.05e 0.029 ± 0.004d 0.21 ± 0.02d 0.41 ± 0.03e 0.012 ± 0.002c
0.05 0.25 ± 0.01d 0.58 ± 0.03d 0.032 ± 0.003d 0.41 ± 0.02c 0.54 ± 0.02d 0.014 ± 0.002bc
0.025 0.42 ± 0.03c 0.89 ± 0.02c 0.089 ± 0.002c 0.48 ± 0.02b 0.76 ± 0.03c 0.016 ± 0.004b
0.0125 0.82 ± 0.03b 3.32 ± 0.03b 0.158 ± 0.006b 0.56 ± 0.02a 1.37 ± 0.04b 0.042 ± 0.002a
CK 0.93 ± 0.02b 3.49 ± 0.04a 0.191 ± 0.004a 0.55 ± 0.01a 1.78 ± 0.01a 0.041 ± 0.002a
  
Leaf 0.1 0.00 ± 0.00d 0.00 ± 0.00d 0.000 ± 0.000d 0.00 ± 0.00d 0.00 ± 0.00e 0.000 ± 0.000e
0.05 0.00 ± 0.00d 0.00 ± 0.00d 0.000 ± 0.000d 0.33 ± 0.03c 0.48 ± 0.03d 0.012 ± 0.001d
0.025 0.37 ± 0.03c 0.84 ± 0.03c 0.075 ± 0.001c 0.46 ± 0.03b 0.71 ± 0.05c 0.018 ± 0.001c
0.0125 0.79 ± 0.03b 3.26 ± 0.03b 0.152 ± 0.007b 0.52 ± 0.04a 1.29 ± 0.04b 0.033 ± 0.003b
CK 0.93 ± 0.02b 3.49 ± 0.04a 0.191 ± 0.004a 0.55 ± 0.01a 1.78 ± 0.01a 0.041 ± 0.002a
  
Flower and fruit head 0.1 0.00 ± 0.00d 0.00 ± 0.00d 0.000 ± 0.000d 0.00 ± 0.00c 0.00 ± 0.00d 0.000 ± 0.000d
0.05 0.00 ± 0.00d 0.00 ± 0.00d 0.000 ± 0.000d 0.00 ± 0.00c 0.00 ± 0.00d 0.000 ± 0.000d
0.025 0.24 ± 0.03c 0.63 ± 0.04c 0.050 ± 0.001c 0.25 ± 0.02b 0.56 ± 0.04c 0.018 ± 0.001c
0.0125 0.67 ± 0.02b 2.70 ± 0.08b 0.135 ± 0.005b 0.53 ± 0.04a 1.10 ± 0.05b 0.033 ± 0.003b
CK 0.93 ± 0.02b 3.49 ± 0.04a 0.191 ± 0.004a 0.55 ± 0.01a 1.78 ± 0.01a 0.041 ± 0.002a
  
F value Plant part 203.87∗∗ 598.54∗∗ 221.54∗∗ 391.59∗∗ 383.81∗∗ 63.88∗∗
Concentration 2650.04∗∗ 33,317.78∗∗ 7095.58∗∗ 1288.43∗∗ 6803.15∗∗ 1127.35∗∗
Interaction 28.46∗∗ 91.84∗∗ 23.49∗∗ 71.36∗∗ 59.60∗∗ 17.93∗∗
  • Note: Means with different letters are significantly different (p < 0.05).

3.3. Biomass of Seedlings

All aqueous extracts, except for the root and stem extracts at 0.0125 g·mL−1 for C. coronarium, demonstrated powerful inhibitory effects on the seedling biomass of both species. The inhibitory rates of the aqueous extracts on the biomass of B. rapa and C. coronarium significantly increased with higher concentrations, with B. rapa experiencing greater inhibition than C. coronarium.

3.4. Allelopathic Response Index

For B. rapa, all measured allelopathic response indices were significantly below zero, except for the root extracts of 0.0125 g·mL−1, which had a positive effect on shoot length. The allelopathic response indices of the aqueous extracts on B. rapa were reduced significantly with increasing concentration. Comparing anatomical sources of the extract, the magnitude of the inhibition was greatest for flower and fruit head extracts generally, with somewhat less inhibition for leaf extracts, and slightly lower levels for stems, and least inhibition by root extracts (Figure 2). Comparing impacts on growth parameters for B. rapa, shoot and root length and biomass were generally more inhibited than germination index, with germination rate the least inhibited (Figure 2).

For C. coronarium, some allelopathic response indices at low concentrations 0.0125–0.025 g·mL−1 were above 0 (i.e., stimulatory); other allelopathic response indices were markedly below 0, however. Allelopathic response indices for C. coronarium were significantly lower with increasing concentration, and flower and fruit head extracts generally were most inhibited and the affected plant parameters followed a similar pattern as seen in B. rapa (Figure 3). Collectively these results showed that the overall inhibition rates on the two bioassay species followed the order flower and fruit head > leaf > stem > root, and that the inhibitory rates for B. rapa were generally higher than those for C. coronarium.

4. Discussion

The invasiveness of many plant species is enhanced by allelopathy [9, 21]. Release of allelochemicals by invasive plants has been shown to inhibit native plants physiologically, even to the extent of causing local extinctions [7, 2224]. Recent studies have indicated that many invasive plants can suppress growth of major crops via allelopathy [3, 6, 2224]. However, with A. radicans​ being a recently recorded invasive species for Yunnan Province, China, its allelopathic effects on neighboring plants, especially major vegetable crops, were largely unknown prior to the current study.

The current study demonstrated various impacts of A. radicans extracts on the early life stages of B. rapa and C. coronarium based on the plant part, concentration, and interaction level. These findings were similar to the patterns seen in some previous studies [2528]. Acmella radicans extracts impacted B. rapa and C. coronarium at their most vulnerable life stages through allelopathy. Root length was most severely inhibited in both species, followed by biomass, shoot length, and germination index, with the germination rate experiencing the least inhibition. The successful establishment of invasive plant species in new habitats often requires high rates of seed germination [22]. Reduced root length, extended germination time, and delayed seedling emergence caused by invasive alien plants can significantly impact how well native plants compete for both aboveground and belowground resources [22]. Methanol extracts from the invasive alien plant Ipomoea cairica delayed the seed germination of two vegetable species, Lactuca sativa and L. sativa var. ramosa, and the roots treated with the extract displayed evident shortness and stoutness, reduction of root hairs or no root hairs, and brownness or death [28]. Allelopathic inhibition of germination and root length often leads to decreased water and nutrient absorption, reducing effective resource utilization and negatively impacting growth, species viability, and ultimately resulting in reduced plant populations [22, 28].

Production of allelochemicals varies greatly in terms of timing and plant anatomical sources [2527]. Allelopathic compounds may be released in various ways, including via volatile compounds, shoot or root leachates, root exudates, and during decomposition in the soil [8]. Numerous studies have indicated that leaf material typically exhibits stronger allelopathic effects and may possess greater concentrations of allelochemical compounds [25, 27]. Fernandez et al. [27] found that the concentrations and diversity of allelochemical compounds were more in the leaves than in roots of Pinus halepensis. The current study found that the inhibition of seed germination and seedling growth of B. rapa and C. coronarium by A. radicans extracts was generally higher for extracts from flowers and fruit heads compared to leaves and stems, with the lowest rates observed for roots. Additionally, the vast majority of allelopathic response indices for A. radicans extracts consisted of negative values that significantly decreased with increasing concentration.

Jirovetz et al. [16] identified more than 70 constituents in the essential oil produced by A. radicans in Southern India. It is difficult to determine which of these are active against plants without further testing. The major compounds they identified were 1-pentadecene, 2-tridecanone, pentadecane, sesquiterpenes, and tridecane [16]. Interestingly, they determined through GC analysis that palmitic acid comprised 16.6% of the volatiles from A. radicans [16]. A study of inhibition of invasive plants by various compounds in sweet potato (Ipomoea batatas) found that palmitic acid produced the second highest levels of inhibition in the 4 invasive alien plants tested [29]. Thus, the allelochemicals produced by A. radicans are likely volatile materials primarily released from aboveground plant parts.

Kato-Noguchi et al. [30] were the first investigators to identify allelopathic activity in the congeneric species, Acmella oleracea, demonstrating inhibition of four plant species by A. oleracea substances. They isolated two substances (E, E)-2,4-undecadien-8,10-diynoic acid isobutylamide and nona-(2Z)-en-6,8-diynoic acid 2-phenylethylamide from A. oleracea responsible for the inhibitory effects [30]. Moreover, the phytotoxic potential of A. oleracea extracts and fractions against three plants such as L. sativa, Calopogonium mucunoides, and Ipomoea purpurea was explored and some allelochemicals were identified through GC/MS analysis [31]. The inhibitory effects of these compounds from A. oleracea may be very useful to determine if these or similar compounds produced by A. radicans also show phytotoxic activity. Some active allelochemicals such as (E,E)-2,4-decadienal, 2-tridecanone, γ-cadinene, δ-cadinene, (E)-α-cadinol, spathulenol, caryophyllene oxide, and widdrol in essential oil from A. radicans were further explored in another study [32].

Acmella radicans has been considered as a noxious invasive alien species in north Yunnan Province, China [17]. In a separate study, the MaxEnt model indicated that in China the potential suitable area for A. radicans is primarily located in southern regions of the country. This plant species has demonstrated strong competitive ability when grown with other plants as it can reduce population densities and importance values of many plant species and absorb more nutrients in invaded habitats [17]. Moreover, the seed germination and seeding growth of some associated weeds were markedly inhibited by aqueous extracts of A. radicans [33]. The strong allelopathic potential of A. radicans against B. rapa and C. coronarium has implications for managing A. radicans, in that it appears the negative impacts are not just due to resource competition. Thus, in order to reduce or avoid the allelopathic effects of A. radicans on crop growth, some measures such as utilizing alternative crop species, variation in crop planting times, and removal of A. radicans plant residues should be considered. In addition, positive uses of A. radicans could be explored such as inhibiting neighboring weed species or developing other ways to make use of its herbicidal properties.

5. Conclusion

The findings of this study demonstrated that aqueous extracts from A. radicans significantly inhibited growth of B. rapa (field mustard) and C. coronarium (garland daisy). The germination rate, germination index, root length, shoot length, and biomass were markedly reduced as the concentration of A. radicans extracts increased. The highest inhibition rates on the two bioassay species were observed for root length and biomass, followed by shoot length and germination index, with the seed germination rate experiencing the least inhibition. Among the allelopathic response indices derived from the aqueous extracts of A. radicans, flowers and fruit heads exhibited the strongest inhibitory effects, followed by leaves and stems, while root extracts had the least impact. Additionally, inhibition by A. radicans was generally greater for B. rapa compared to C. coronarium. Most allelopathic response indices for aqueous extracts of A. radicans significantly decreased with increasing concentration, providing strong evidence of its allelopathic inhibition. To gain a more comprehensive understanding of the allelopathic mechanisms of A. radicans, additional investigations are required to identify its allelochemicals and elucidate the biochemical mechanisms involved in its allelopathic activity.

Conflicts of Interest

The authors declare no conflicts of interest.

Author Contributions

S.S. and F.Z. conceived and designed the experiments; G.X., S.Y., G.J., K.Y., X.W., and Z.F. performed the experiments; S.S. and D.R.C. wrote the draft. The final manuscript was read and approved by all authors.

Funding

This research was supported by the National Key R&D Program of China (2021YFC2600400), Ten Thousand Talent Program (Young Top-Notch Talent) of Yunnan Province (YNWR-QNBJ-2018-201), and Key Research and Development Program of Yunnan Province (202103AF140007, 202203AE140008, and 2019IB007).

Acknowledgments

We are grateful for Randi Wu from the Agriculture and Life Sciences College of Kunming University and Yuchen Cui from the School of Agriculture, Yunnan University, for expert technical assistance.

    Open Research

    This published article incorporates all the data we collected and analyzed.

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