1. Introduction
Canola (
Brassica napus L) is an oil seed crop belonging to the Brassicaceae family. It results from the natural hybridisation between
Brassica oleracea L. and
Brassica rapa L. Australia is the world’s second largest exporter of canola [
2]. Canola grown in Australia meets the high expectations of exporters, domestic crushers and intensive livestock producers. However, weeds are an important biotic constraint on canola production, resulting in yield and quality losses [
4,
45]. Grass weeds such as annual ryegrass (
Lolium rigidum), vulpia (
Vulpia myuros) and wild oat (
Avena fatua) are the most abundant weed species in canola crops of south-eastern Australia [
30]. Weeds from Brassicaceae can contaminate canola seed samples which leads to increased levels of erucic acid and glucosinolates with the consequent reduction in canola quality due to their contamination [
4,
45].
Chemical herbicides and mechanical weeding have been the most frequently used methods to control weeds [
16,
26] and have served to keep weed infestations low, thereby improving crop productivity. Despite the significant contribution of these methods, there are also certain associated challenges. Mechanical weed control requires extra soil turn-over, which can disturb soil structure and deplete soil fertility [
48]. Additionally, it is not always effective and can be expensive and lack durability [
15]. Similarly, herbicide-resistant weeds, health effects and environmental concerns are the major constraints for repeated use of herbicides [
7,
49]. In Australia, non-chemical weed control options are highly sought after for incorporation into improved integrated weed management systems (IWMS) to overcome herbicide-resistance.
Crop plant interference against weeds involves the combined effects of plant competition and allelopathy. Competition is the negative interaction between two or more plant species for resources within a limited space and is distinct from plant allelopathic interaction [
56]. The competitiveness of a plant is influenced by various morphological, phenological and agronomic traits [
25,
26,
31,
32]. Increasing crop seeding rates is one of the simplest agronomic ways to enhance crop competitiveness [
35], as it increases crop canopy development and hastens nutrient use, thereby denying these resources to weeds and thus reducing weed pressure [
10,
51]. In contrast to competition, allelopathy is the exudation of compounds by plant roots that can suppress the growth of neighbouring plants and affects seeds and seedlings of other species located within a limited range [
39]. Although most plants species, including crops, can produce and release biologically active root exudates (allelochemicals), relatively few have strong allelopathic properties. Several bioactive compounds were isolated from strongly-allelopathic canola genotypes [
3]. These phytotoxic or signalling chemicals presumably resulted in the observed inhibitory effects on annual ryegrass (
L. rigidum) under laboratory conditions, and also may be responsible for the significant suppression of other weed species in the field [
5]. Allelochemical concentrations are a function of the density of the allelopathic crop [
9]. This suggests that density may be an important factor in enhancing canola allelopathic activity.
Crop species and genotypes within the species differ in root traits and their ability to compete for below-ground resources, which in turn can influence aboveground traits and yield [
17,
55]. Root and shoot architecture including length, biomass, number of lateral roots and growth play a crucial role in competition and crop performance [
6,
18,
23,
41]. Hence, screening crop varieties with plant functional traits may offer new insights into plant allelopathy. Further, plants can perceive different external and internal signals from their surroundings, while changes of environmental conditions subsequently can affect plant allelopathy and its functional traits [
40]. Huang et al. [
28] demonstrated that
Merremia boisiana can adjust its resource allocation to allelopathy and leaf functional traits to adapt to varying environments. As one of the key factors affecting plant allelopathy and the functional traits, environmental changes have long been assumed critical. However, the link between canola allelopathy and its functional traits has received little attention.
Understanding how crop genotypes respond differently to changing environmental conditions is crucial and it is a significant step in developing improved crop varieties [
43]. When genotypes are assessed across various locations or years, their yield and individual traits’ performances may vary significantly. The presence of substantial genotype-by-environment (G × E) interaction can further complicate comparisons and recommendations for adaptable genotypes [
36]. To identify stable genotypes, it is essential to break down G × E interaction into stability statistics assigned to each genotype across different environments. Various stability indices have enabled researchers to pinpoint widely adapted genotypes for breeding programs or enhance recommendations to growers [
38]. A genotype is considered most stable when it exhibits minimal fluctuation across diverse environments [
38]. The present research aimed to examine 1) the functional traits and weed suppressive ability of four selected canola genotypes and 2) the stability of canola genotypes for weed suppression over different temporal and spatial conditions.
4. Discussions
The study of plant interference is increasingly popular although the current systems of plant breeding largely ignore the abilities of a variety to exercise control over its weed challengers. The difficulty in studying interactions between plants is due to the complex nature of plant interference, defined as the combined effect of competition and allelopathy. However, the development of crops with the capability to exert allelopathic effects on weeds through root exudates is an attractive prospect [37. Research has shown this potential in wheat [
53], barley [
11,
12], rice [
19,
20,
21,
24,
46] and canola [
5]. In this study, genotypes such as AV-OPAL, PAK85388-502, AV-GARNET and BAROSSA produced similar crop biomass but the genotypes PAK85388-502 and AV-OPAL tended to result in weed biomass lower than even the strongly competitive genotype (cv. AV-GARNET). Therefore, weed-suppressing genotypes have potential for integrated weed management and further work is needed to produce a genotype with combined competitive ability and allelopathy. Bertholdsson [
12] found that early weed biomass was significantly lower in the highly allelopathic wheat lines compared with the non-allelopathic lines. Also, our research revealed that weed biomass was suppressed during the early growth stages of allelopathic canola genotypes, resulting in significantly less weed biomass harvested at the end of the experiment.
Four Brassica genotypes (AV-OPAL, AV-GARNET, BAROSSA PAK85388-502) showed similar patterns in the density-response curve for weed growth and there was a density by genotype interaction. This indicates that crop density plays a role in canola weed suppression. These results were consistent with previous research in rice [
46] and in wheat [
36]. The aboveground canopy of all tested genotypes had similar contributions to weed control. However, beyond that, the additional mechanism such as the allelopathy of AV-OPAL and PAK85388-502 might played a role for their neighbouring weed suppression. AV-OPAL was identified as a less vigorous genotype with shorter plant height among other canola genotypes [
5]. Asaduzzaman et al. [
6] collected 70 international rapeseed varieties and evaluated their allelopathic potential by growing them in close proximity to
L. rigidum. Rapeseed was sown at 10, 20, and 30 plants/m
2 against 15 plants/m
2 of annual ryegrass. Generally, the higher density of rapeseed resulted in higher suppression of
L. rigidum. The varieties with strong allelopathic activity were PAK85388-502, AV-OPAL, BLN3343CO0402, and RIVETTE. Many of these genotypes categorised as allelopathic those can released a range of allelochemicals through their root exudates of canola, with key chemicals being sinapyl alcohol, p-hydroxybenzoic acid and 3,5,6,7,8-pentahydroxy flavones [
3]. Furthermore, in Canada, canola competitiveness was improved by choice of variety and use of higher seeding rates [
8]. Correlation coefficient is a measure of the extent and direction of the relationship between any two traits (variables). The negative relationships between root (number of lateral roots) and shoot (stem density, and stem length) functional traits and weed biomass suggest that these traits are important components for weed suppression by canola genotypes. Therefore, their simultaneous selection will be a good approach to increasing weed suppressive ability. This same relationship was observed by Afuape et al. [
1]. Genetic variability is essential for selection [
44]. However, a wide range of varieties should be evaluated at different locations to further verify such relationships.
The stability analysis aims at helping breeders identify which genotypes have specific and/or general adaptability to various production environments. Additionally, stability analysis helps determine the test environments for future evaluations of canola production with self-weeding capability. Three locations (excluding Wagga 2013) clustered in the same quadrants, indicating that these locations share similarities in terms of weed growth and canola genotype performance. The Temora environment and Wagga 2016 were higher performing environments for weed suppression compared with others. This result means that testing data from one location can represent the performance of same materials in the other similar locations. Therefore, conducting a stability analysis with a large set of canola genotypes will further help identify specific genotypes for both locations, as well as a stable genotype that can be cultivated across multiple locations and particularly locations that share similar attributes to the test locations. Understanding the environmental conditions in which canola varieties are tested is crucial for accurately interpreting results and extrapolating findings to diverse geographical locations or growing conditions. Highlighting higher-performing environments for weed suppression, such as the Temora environment and Wagga 2016, underscores the need for a comprehensive description of these locations' environmental characteristics. To address this gap, future research should prioritise conducting stability analyses with a diverse range of canola genotypes across various locations. This approach would enable breeders to identify specific genotypes suited for different environments and stable genotypes adaptable to multiple locations, facilitating informed decision-making in canola allelopathic variety cultivation.
The weed suppressive ability of a specific canola variety might be weed-specific and likely associated with a genetic as well as an environmental component of variance; breeding will be required for its maintenance [
11]. Combining a variety’s capability in reducing specific weed pressure with optimal agronomic practices that facilitate crop health will generally enhance cropping system sustainability and allow growers to extend the life of valuable herbicides [
27]. However, the feasibility of using a variety for specific weed management may not be useful for economic outcomes because canola crops are infested naturally by a range of weed species [
30,
33]. However, a competitive variety of canola could be developed by incorporating traits such as increased lateral roots, stem length and density without compromising other desirable traits such as grain yield, quality or disease resistance [
31]. Canola weed suppressiveness can be optimised to reduce specific weed growth and reproduction through farming practices that allow implementation of a variety of cultural techniques such as sowing crops with different planting dates to reduce other weeds pressure [
51]. In addition, the ability of crops to suppress weeds appears to be strongly variety dependent [14; 29; 54).
Here, the experiments reported reinforce the need for the preservation of the older varieties such as AV-OPAL and PAK85388-502 (highly allelopathic) and AV-GARNET (highly competitive [
31]) so that these benefits can be incorporated into new varieties. Also, this research highlights the need for new varieties to be evaluated for their interference capabilities in weedy field plots without herbicides. The information obtained in this future research will allow producers to broaden their armoury against herbicide resistance by choosing weed-suppressive varieties, thereby helping manage herbicide-resistant weeds which threaten productivity, profitability and food-security. The capability of crop variety to suppress weeds is being considered as a preferred criterion for cultivar selection in many parts of the world [
52]. The allelopathic potential of crop plants contributes to the weed suppressing ability of the variety. Weed-suppressive allelopathic varieties can reduce weed infestation and improve the efficacy of inputs and increase the methods of weed control.