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Article

The Relationship Between the Germination of Silky Bent Grass (Apera spica-venti (L.) Beauv.) Diaspores and Their Age, Place of Occurrence, and Action of Stimulating Substances

Institute of Agroecology and Plant Production, Wrocław University of Environmental and Life Sciences, Grunwaldzki Sq. 24A, 53-363 Wrocław, Poland
Agronomy 2025, 15(3), 715; https://doi.org/10.3390/agronomy15030715
Submission received: 6 February 2025 / Revised: 6 March 2025 / Accepted: 13 March 2025 / Published: 15 March 2025

Abstract

:
Apera spica-venti is a weed that is threatening agricultural crops worldwide. Current reports do not fully agree on the biology of the weed, regarding the viability of diaspores, nor is there sufficient information on how different factors affect the germination of its seeds, such as the abundance of soil where the mother plant (plants from which diaspores (caryopsis) were collected) has grown or the effect of biostimulants on their germination. Therefore, the main aim of this study was to compare the germination of newly harvested seeds of A. spica-venti (3 months after harvest) with seeds that had been stored for 1, 2, 3, 4, or 5 years. Furthermore, the influence of seed age, weed location, and biostimulants (silicon and algal liquids) on the germination of A. spica-venti diaspores was checked. Three-year-old seeds showed the highest percentage of germination, and their germination process was positively correlated with soil abundance (from sites of mother plant material collection) and macronutrients (N, P, K). The strongest correlations were recorded for 3- and 4-year-old seeds. A. spica-venti seeds treated with biostimulant preparations were characterized by varying percentages of germination. No clear trend was noted regarding the germination capacity of the harvested seeds as the diaspores aged, and it depended on the type of stimulant and the place of origin of the seed. Moreover, seeds from the harvest year treated with the stimulant were characterized by a higher germination percentage. A. spica-venti is a highly fecund weed, a trait that makes it very common in crops, appearing en masse and providing formidable competition to crops, thus causing significant yield losses. Awareness of the vitality of diaspores will allow for the possible regulation and control of this weed in order to prevent yield loss in crops. The theme of diaspore viability warrants further investigation. Further research should include observations of factors affecting germination, including new stimulants emerging on the agricultural market.

1. Introduction

Noxious weed silky bent-grass (Apera spica-venti (Linnaeus) P. Beauvois) occurs commonly in crops regardless of the intensity of farming and the climate zone [1,2]. A. spica-venti is a highly fecund weed. As reported by [3], one plant is capable of producing approximately 2000 to 20,000 diaspores, depending on crop competition. Various definitions of the term diaspore can be found in the literature. It is most commonly defined as both generative, such as spores, seeds or fruit, and vegetative, such as fragments of stems, stolons, rhizomes, or bulbs [4]. Alternatively, the term ‘diaspore’ (or ‘dispersule’) generally refers to the dispersal unit, regardless of which part of the plant this definition applies to [5]. Diaspores of the weed are the of orthodox type [6]. They germinate in a wide temperature range and in limited light conditions, which explains their more rapid growth compared to arable crops. As a consequence, A. spica-venti spreads rapidly, especially in winter cereals [7].
In the opinion of Nautiyal et al. (2023) [8], germination can be defined in physiological terms as the metabolic activation of the seed upon hydration, culminating in ‘chitting’ or protrusion of the radicle through the seed coat. According to reports, many factors influence the germination of weed seeds/diaspores, starting with the genetic conditions of the species [9,10], the growth and development conditions of the mother plants (soil nutrient abundance, periods of drought) [11], the age of the seeds [12], maturation of seed [13], seed storage conditions [14], and their stage of dormancy [15].
Furthermore, the dormancy of seeds is conditioned by the course of metabolism, which is genetically determined by protein synthesis or allosteric variability of enzyme activity. The ability of weed diaspores to enter dormancy determines their longevity [16]. However, according to Bochenek [17] and Bochenek et al. [18], the length of dormancy can be modified by the influence of external factors on the seed coat. Grzesiuk and Łuczyńska [19] suggested that the dormancy of grass seeds is caused by the impermeability of the seed coat to gasses. This type of dormancy occurs due to the inhibition of enzyme action by growth inhibitors [19]. Additionally, Baskin and Baskin (1998) [20] point out that species growing in environments with high levels of competition produce seeds with the highest germination rates to remain competitive with other species, which is very consistent with the characteristics of the species under study.
Plants of A. spica-venti that emerge in autumn have higher vitality and form more populous stands than those emerging in spring [3]. The viability of A. spica-venti diaspores is still the subject of debate. However, diaspores of this species can retain their vitality over a period of 1–2 years or for as long as 7 years [21,22,23]. Kukowski [24] showed the germination capacity of diaspores stored under laboratory conditions, respectively, after 3 months at 55%, up to 2 years at 33%, and in the 3rd year at 48%. In subsequent years, a decrease in germination was noted, with a capacity of 6% after 5.5 years. However, Hanelt [25] proved that diaspores stored at room temperature can remain viable for up to 11 years, with a germination capacity of 9%.
The economic losses caused by a given weed and its common occurrence make it the subject of research on its resistance to active substances contained in herbicides [26,27,28] and the effects of these substances on crop yields [29]. Against the background of a warming climate, along with other climatic changes, plants are subjected to increasing stress levels [30,31]. As a consequence, agricultural producers frequently use biostimulants to limit the negative impacts of abiotic factors on crops [32,33,34]. The substances contained in these preparations are intended to increase the resistance of plants under stress conditions and act as stimulants of life processes [35].
The name “biostimulant” refers to a large group of products that differ in terms of their contents of active substances. They can contain substances of natural origin, microorganisms, and synthetic chemical compounds as active substances [36,37]. The effects of these preparations on the cultivation of agricultural plants depend on the type of crop and composition [38]. These preparations can be applied as alternatives to chemical formulations to improve crop yield [39,40]. Biostimulants include, among others, preparations based on marine algae such as Ecklonia maxima and Ascophyllum nodosum [41], or those containing a specific element [36,37]. According to report [36], seaweed extracts are marketed as biostimulants because they contain multiple growth regulators such as cytokinins [42], auxins [43], gibberellins [44], and various macro- and micronutrients necessary for plant growth and development. They also promote tolerance to environmental stress [45]. According to Mansour et al. [46] and Matysiak and Miziniak [47], preparations based on algae can increase the yields of winter wheat [46] and oilseed rape [47] in areas where A. spica-venti commonly occurs. Biostimulants can contain sea algae extracts, free amino acids, and humic compounds, along with nutrients, including silicon (the so-called “synthetic biostimulators”). Silicon availability to plants is generally limited because it mainly occurs in the form of silica (SiO2), which is insoluble in the soil solution. Silicon-fertilized plants are characterized by increased resistance to disease and salinity [48].
Generally, preparations that contain silica reduce transpiration and the sensitivity of plants to light deficiency [49]. In addition, they can also cause endodermal silification, protecting plant roots from pathogens (this is important for cereals, whose main fungal pathogens are Fusarium fungi, which are so-called “soil-borne phytopathogens”) and parasites. The above advantages of silicon increase plant yield and improve yield quality [50]. Because cereal plants and grasses can accumulate up to 3% silica whereas dicotyledons accumulate less than 0.5% [40], the use of preparations containing silicon can increase the resistance of A. spica-venti to stress, prolonging the lifespan of the seeds [51].
As indicated by Gupta et al. [36], the germination of seeds (heterotrophic phase) and seedling growth (autotrophic phase) is the most crucial and sensitive stage in the life cycle of a plant. Although it has been reported that biostimulants based on algal extracts can stimulate seed germination due to the content of phytohormones [52], they may also have an inhibitory effect on the process due to their possible salt content [53]. Reports on the effect or non-effect of biostimulants mainly concern crop plants [52,53]. Weeds are companion plants of agricultural crops, and therefore their response to the applied biopreparation cannot be ignored. Currently, there are no reports in the literature on the effects of biostimulants based on algae or silicon on weed diaspores.
The main purpose of this study was to determine whether there is a difference in seed germination between newly harvested seeds of A. spica-venti (3 months after harvest) and seeds that had been stored for 1-, 2-, 3-, 4-, or 5 years. Additional objectives included checking (1) whether the harvest site influences seed germination, (2) how seed germination changes depending on age, and (3) the influence of biostimulants such silicon as liquids (SiO2) and algal liquids from E. maxima on the germination of A. spica-venti diaspores.

2. Materials and Methods

This study was conducted in the Institute of Agroecology and Plant Production, Wrocław University of Environmental and Life Sciences and Sciences, Poland, from 2016 to 2021.

2.1. Weed Diaspores

The tested weed diaspores of A. spica-venti was sampled randomly from crops located in the Lower Silesia Province (Poland) (Table 1, Figure 1).
Plant material (diaspores) was collected in the same areas over the years 2016–2019. This weed did not occur in the study area in each year of the study period (weed harvesting was carried out on cultivated fields).
Seeds to research were collected from various crop fields (Table 1) after maturation of the seed. The mature weed plants were harvested on 8–12 July 2016. In 2017, the weed harvest date ranged from 6 to 18 July 2017. In 2019, the weed was harvested from 2 to 3 July. Seeds were put into paper bags to keep them dry and to avoid humidity and climatic factors which lead to germination; they were kept under laboratory conditions (room temperature 21–22 °C, humidity 45–50%). Information regarding soil chemical analyses—Table A1—Appendix A: The Influence of the Habitat on the Chemical Composition and Morphology of Silky Bent Grass (Apera spica-venti (L.) Beauv.) Occurring in Arable Fields (Lower Silesia, Poland). Agronomy 2022, 12, 1883. https://doi.org/10.3390/agronomy12081883 [54].

2.1.1. A. spica-venti Collection Region

Lower Silesia is located in south-western Poland and borders Germany and the Czech Republic. A characteristic feature of the region is the great diversity of its relief and landscape. This is due to the fact that several geographical regions, including lowlands and foothills, are located in the area. The occurrence of different physiographic conditions results in significant climatic variations. The northern part of the region (where plant material was collected) is located in the moderate zone of the oceanic and continental climates. The southern part of the region is under the influence of a zonal mountain climate. The northern part of the region is one of the warmest areas in Poland. The annual precipitation is 550–700 mm and the average annual temperature is around 8 °C. The vegetation period in the area is 220 days.

2.1.2. Global and National Distribution of the Weed

A. spica-venti is the most common grass weed species in winter cereals in Denmark, Germany, Poland, Czech Republic, Slovakia, Lithuania, Latvia, and Austria, but also present in Belgium, The Netherlands. Luxembourg, France, Switzerland, Sweden, Belarus, Ukraine, and Russia [3]. According to reports, the areas where the weed occurs are characterized by a higher proportion of winter annual crops in rotation [3].

2.1.3. Taxonomy of the Weed

Apera spica-venti (Linnaeus) P. Beauvois belongs to class Equisetopsida, subclass Magnoliidae, superorder Lilianae, order Poales, family Poaceae, subfamily Pooideae, tribe Poeae, subtribe Poinae, and genus Apera species Apera spica-venti [55].

2.1.4. Diaspores Description of A. spica-venti

The diaspore traits of A. spica-venti are characterized by the following parameters: shape of caryopsis—oblong, longitudinally grooved on the ventral surface, normal size—1.0 to 1.5 mm long by approximately 0.5 mm wide; color—reddish-yellow; texture—smooth and glabrous; style base usually remaining attached at apex; embryo—approximately one-third the length of the caryopsis; endosperm—soft to semi-liquid; hilum—basal, ovate [56].

2.2. In Vitro Experiment in Petri Dishes

Germination analyses were carried out in a commercial vegetation chamber [MLR-352H/SANYO/Moriguchi/Japan] during 2016, 2017, 2019, 2020, and 2021 always after the seeds had reached the required age (counted from the date of harvest). The individual test performed in glass Petri dishes containing two layers of tissue paper and sterilized at 130 °C. The filter paper were soaked under aseptic conditions with 10 mL sterile distilled water or of the preparation, depending on the test variant (Table 2).
A total of 100 diaspores of A. spica-venti from each location (Table 1) placed on the Petri dishes in four replicates, followed by incubation in a plant chamber for 30 days. During this period, A. spica-venti germinated seeds were counted after 5, 10, 15, 20, and 30 days. Additionally, after 30 days, leaf length was determined. Life span assessments were carried out in accordance with the planned life testing option (Table 2). The tested active substances recommended to stimulate resistance to external factors were liquids of silicon and algae. Silicon was in the form of SiO2 (200 g·L−1)—the commercial product (OPTYSIL®) was used (0.5 L·ha−1 for winter wheat spraying in one time, recommended number of treatments 4). The second active substance tested was a liquid concentrate made from Ecklonia maxima. The commercial product (Kelpak SL) was also used (2 L·ha−1 for winter wheat spraying in one time, recommended number of treatments 1). The concentrations of auxins and cytokinins is 11:0.031 mg·L−1. However, the active ingredient was diluted 4 times, due to the fact that diaspores stagnated in solution. The conditions in the vegetation chamber were planned to take into account some the suggestions by Morales et al. [57], where the authors, in their conclusions, suggest certain settings for seed germination parameters. In their preliminary test, like photoperiod 14 h of daylight and 10 h of darkness. Seeds in their experiment were not pre-chilled and no KNO3 was applied. As the authors themselves suggest, the knowledge about seed germination is negligible [57].

2.3. In the Study Determined Indicators

Final Germination Percentage (FGP) (%)—Number of total seeds germinated over the number of total seeds sown time [58].
Dynamics of germination: seed germination was measured after 5,10, 15, 20, and 30 days counted. Dynamics were presented for A. spica-venti seeds from the 2016, 2017, and 2019 harvest years depending on the harvest site.

2.4. Statistical Analyses

The data obtained from this study were analyzed using the Statitica 13.6 package (StatSoft Polska sp. z o.o., Kraków, Poland). For this purpose, one-way analysis of variance (ANOVA) and Tukey’s HSD (honest significant difference) test at α ≤ 0.05.
Prior to the ANOVA, the percentage data were transformed to Bliss [59] angular degrees by applying the formula y = arcsin (value%)−0.5. After transformation, the variance was approximately constant, allowing the ANOVA to compare particular components [59].
The Pearson (r) correlation coefficient at α = 0.05 was used to determine the relation between the mean levels of N, P, and K in the soil and pH of soil with diaspores germination.

3. Results

3.1. Germination of A. spica-vention

Statistical analysis of the obtained results showed significant variation in the percentage of germinated diaspores depending on the site of occurrence (Table 3). For seeds from a harvest year, the difference between the best and worst germination seeds was 22.5 percentage points. In addition, for seeds that were several years old, the difference ranged from 16.0 (3-year) to 24.5 (2-year) percentage points. In the case of some diaspores, it was found that, depending on the site of occurrence, the best values in terms of germination percentage were reached at the age of 2 or 3 years (respectively 56.25 and 55.75%).

3.2. Age of Diaspores and Germination

Significant differences in seed germination were found depending on the age of the diaspores independently for a harvest site. Seed viability varied with age. The highest value was found in 3-year-old diaspores (43.55%) (Figure 2). The percentage of germination of 3-year-old seeds was significantly different from the value achieved by other diaspores (for this year’s, 1-year, 2-year, 4-year, and 5-year diaspores p = 0.000020; p = 0.000054; p = 0.002253; p = 0.000287; p = 0.000021, respectively).
A similar percentage of germination was found for 1-year-old and 4-year-old diaspores (23.55% and 25.4%) (Figure 2). The recorded values were significantly lower than the highest recorded value (3-year diaspores). On the other hand, 5-year-old diaspores and diaspores germinated in the year of harvest were characterized by similar values (13.25 and 14.97%, respectively).

3.3. Dynamics of Germination

Analysis of the germination dynamics of A. spica-venti diaspores showed significant differences depending on their age and site of occurrence. However, this was not clear in all analyses. The highest changes in seed germination were recorded, depending on the collection site and age gain during the 15 days of observation.
The discrepancies in the analysis of the germination dynamics of diaspores differing in age may indicate the conditions that occurred during the period of growth and development of the mother plant, presumably depending on the abundance of the soil in the site of occurrence and the weather (Figures S1–S10).

3.4. Correlations of Germination with Macronutrients

The analysis of the results showed a negative weak correlation between the germination of A. spica-venti seeds in the harvest year and nitrogen content in the soil (p < 0.05; r = −0.25) (Figure S11). One-year diaspores were characterized by a positive weak correlation (p < 0.05; r = 0.28) (Figure S12), but weak negative correlation was characterized by 2-year-old seeds (p < 0.05; r = −0.42) (Figure S13) in relation to the nitrogen content in the soil during the harvest period (Table A1 [54] and Table 4). Different results were obtained as a result of the germination analysis of 3- and 4-year-old seeds, where the results were moderate and fairly strongly positively correlated, respectively, p < 0.05; r = 0.69 and p < 0.05; r = 0.72 (Figures S14 and S15).
The correlation values between the germination of seeds from harvest year, 1-year and 2-year diaspores, and the soil’s phosphorus content in the soil were characterized by weak correlation and the absence of a linear relationship; p < 0.05; r = 0.26; r = −0.28; r = 0.05 (Table 4 and Figures S16–S18).
In this research, there was a moderately and strongly positive correlation between the percentage of germinated 3- and 4-year-old diaspores and the phosphorus content in the soil. The recorded values were, respectively, p < 0.05; r = 0.41; r = 0.44 (Figures S19 and S20).
Similar correlations were found for diaspore germination and potassium content in the soil. In the year of harvest, diaspores and one-year old diaspores were characterized by the absence of a linear relationship and a weak negative relationship (p < 0.05; r = −0.15; r = −0.23 (Figures S21 and S22). The 2-year diaspores were characterized by a weak positive linear relationship (p < 0.05; r = 0.20) (Figure S23). In contrast, 3- and 4-year diaspores were characterized by a positive moderate and strong linear relationship, respectively, p < 0.05; r = 0.67; r = 0.95 (Figures S24 and S25). There was no apparent trend between soil pH and germination of A. spica-venti diaspores with aging. The recorded relationships varied from values where no linear relationship noted to a moderate relationship (Table 4 and Figures S26–S30).

3.5. Effect of Biostimulants on Germination

Analysis of the results showed a positive significant effect of biostimulants on the germination of A. spica-venti diaspores in the year of harvest (Table 5 and Table 6). However, this was conditioned by the substance tested. Moreover, changes in seed germination under the influence of the substances tested were observed depending on the age of the diaspores.

3.5.1. Effect of Biostimulants on Germination—Alge

The positive effect of algae-based solutions was recorded for germination of all diaspores in the harvest year from all sites of weed occurrence, with three sites of five being statistically significant (Table 5). In the following years of study, one-year diaspores irrespective of where the plant material was harvested and had a significantly lower percentage of germination compared to control objects (distilled water). A percentage increase in germination was also recorded; however, this concerned diaspores from mother plants from single sites and was not statistically significant. In the case of 2-year-old diaspores from three of five sites of occurrences of A. spica-venti, the germination percentage was significantly higher compared to diaspores germination where the distilled water was used (Table 5). The highest recorded percentage of seed germination was 48.75%. The result obtained was the highest for the algal solution tested over all the years of observation.
The site of appearance of the mother plant significantly differentiated germination of A. spica-venti diaspores. This differentiation was further highlighted by the use of an algal solution.

3.5.2. Effect of Biostimulants on Germination—Silicon

Analysis of the results showed a positive significant effect of biostimulants on the germination of A. spica-venti.
Furthermore, it was noted that there was significant variation in the effect of the silicon preparation on diaspore germination depending on the place of emergence and development of the mother plant (Table 6). The highest recorded percentage of diaspore germination was 52%. The difference between the highest germination percentage and the lowest according to the place of appearance of the mother plant, where the diaspores in Petri dish were flooded with silicon solution, was 39.5 percentage points.
In the case of annual diaspores, only a significant effect of silicon on diaspores from one site was recorded. In addition, an inhibitory effect of a substance was recorded for diaspores from two different mother plant occurrence sites. In case of 2-year-old diaspores, they were characterized by a significantly higher germination percentage under the influence of the test substance. Comparable values in the germination percentage of the diaspores can also be observed for the 2-year-old diaspores and the diaspores in the year of harvest.

3.6. Correlations of Germination with Different Solution—Alge

Correlation analysis of the germination percentage of A. spica-venti diaspores and the macronutrient content showed strong variations according to the age of the diaspores, where the solution used to fill the dishes was the algal solution (Table 7). There was a moderately negative relationship between the germination percentage of diaspores from the year of harvest and the 1-year diaspores and N content in the soil (accordingly p < 0.05, r = −0.45, r = −0.61 (accordingly Figures S31 and S32). The 2-year diaspores showed a weak correlation between the investigated features (p < 0.05; r = −0.29) (Table 7, Figure S33).
In the case of phosphorus, for the germination of diaspores from the year of harvest and annual diaspores, no significant correlation was recorded between their germination with the addition of the algal solution and soil phosphorus abundance (Table 7, Figures S34 and S35). However, there was a moderate relationship between its content in the soil from the place of growth and development of the mother plant and the germination percentage of the 2-year diaspores (p < 0.05; r = −0.64) (Table 7, Figure S36).
However, in the case of potassium content in the soil, the correlations between the traits studied showed a moderate relationship regardless of the age of the seed (in the year of harvest p < 0.05; r = −0.56, 1-year p < 0.05; r = −0.62, 2-year p < 0.05; r = −0.70 (Table 7, Figures S37–S39, respectively). Correlation analysis showed that there was no linear relationship between soil pH from the mother plant site of growth and development and germination of A. spica-venti diaspores (Table 7, Figures S40–S42, respectively).

3.7. Correlations of Germination with Different Solution—Silicon

Person correlation analysis showed variable values for the strength of the correlation between the percentage of diaspore germination depending on age and substance used and the content of NPK components in the soil from site where the plant material was collected (Table 8). The correlation value between the germination of diaspores from the year of harvest was p < 0.05; r = −0.32 (Table 8, Figure S43). One-year-old diaspores were characterized by a very strong negative correlation between their germination percentage and soil N content (p < 0.05; r = −0.97) (Table 8, Figure S44) and two-year and N was r = −0.55, respectively (Table 8, Figure S45), where the substance used to fill the Petri dishes was a substance containing silicon.
The correlation value between the germination of diaspores differing in age and soil phosphorus abundance varied widely. For harvest year diaspores and 1-year diaspores, the correlation between the germination percentage of the diaspores and the abundance of the element under study was p < 0.05; r = −0.15, r = −0.17, respectively (Table 8, Figures S46 and S47). However, for 2-year diaspores, a negative moderate value was recorded (p < 0.05; r = −0.58) (Table 8, Figure S48).
Analysis of the results showed a negative correlation between A. spica-venti seed germination and soil potassium content in the soil. The recorded correlation became stronger as the diaspores gained age. For seeds from the year of harvest, the correlation was p < 0.05; r = −0.52, for 1-year-old seeds p < 0.05; r = −0.62 while for 2-year-old diaspores p < 0.05; r = −0.79 (Table 8, Figures S49–S51, respectively).
The germination percentage of seeds from the year of harvest did not correlate with soil pH p < 0.05; r = −0.08 (Table 8, Figure S52). One-year diaspores correlated quite strongly with soil pH from the place of growth and development of the mother plant p < 0.05; r = −0.71 (Table 8, Figure S53). For 2-year seeds, respectively, it was p < 0.05; r = −0.28 (Table 8, Figure S54).

3.8. Leaf Length

Statistical analysis showed no variation in 1-leaf length within a species differing in the location of the mother plant (2019) (Table 9). Weed seedlings from 1-year-old diaspores also did not differ significantly in the length of the observed leaves. Variable lengths, instead, were recorded for the length of 1-leaf of seedlings originating from 2-year-old weed diaspores. The longest leaves were those of seedlings originating from the mother plant from rye monoculture.
Analysis showed variations in the length of the first leaf of A. spica-venti seedlings depending on the origin of the mother plant at the time of silicon application, in all years of observation regardless of the age of the seed analyzed (Table 10).
Weed seedlings from diaspores in the year of harvest of the mother plant had the longest leaves regardless of the place of their appearance.
Statistical analysis showed differences in the length of the first leaf of A. spica-venti depending on the occurrence of the mother plant under the influence of the algae-based preparation (Table 11).
The lowest values of leaf blade length were recorded for seedlings from annual diaspores, regardless of the place of origin of the mother plant. There was no influence of the tested preparations on leaf blade length elongation of A. spica-venti seedlings in the year of diaspore collection (Table 12).
In the four sites observed, the differences were non-significant, irrespective of the place of occurrence of the mother plant. Only in one case, there was a significant effect of silicon on seedling leaf blade elongation. Seedlings grown from 1-year-old seed had the longest leaves when no silicon or algal preparation was applied. In contrast, the opposite trend was found for seedlings derived from 2-year-old seeds. The seedlings treated with silicon or algal preparation had the longest leaf blades. In addition, there were no differences in the respective years, except for one site of the respective trend.

4. Discussion

The process of seed germination is conditioned by many factors, including light, temperature, ethylene content [60,61], seed dormancy location, seed depth in soil, and the place where the mother plant grew and developed [62,63,64]. The germination of diaspores of a given weed also depends on the conditions created for the seed material’s storage [24]. As Kukowski [24] reported, this is a decisive factor in drawing conclusions about the viability of A. spica-venti seeds. This was also confirmed by other reports [14,65,66]. However, all of them refer to seeds/diaspores. Generally, there is no information on A. spica-vegenerally spores. Those that have appeared do not refer to all storage conditions [24,25].
The analysis of the results of seed germination in this experiment showed that after a 3-month period of dormancy, seeds were put into paper bags to keep them dry and to avoid humidity and climatic factors that lead to seed germination. The seeds were kept in laboratory condition room temperature of 21–22 °C, with a humidity of 45–50%. Seed germination was at the level of 5.75% to 28.25%, depending on the place of harvesting, the plant material, and the year of observation. In the following years of observation (in the 2nd and 3rd years), germination increased, with peak values reached for seeds from harvesting sites, followed by a gradual decrease. Observations showed that 4- and 5-year-old seeds germinated between 9.25% and 20.5%. This is approximately consistent with the results of Hanelt [25] who found that A. spica-venti seeds stored under room conditions maintain viability for up to 11 years. This allows us to speculate that seeds from our own observation data, with a higher percentage of germination, may have 9% viability at 11 years of age. However, this is only speculation as the storage conditions in Hanelt’s [25] study are unknown. Furthermore, Kukowski [24] showed the germination capacity of diaspores stored in laboratory conditions (the author also did not specify the conditions under which the diaspores were stored). After 3 months, the capacity was 55%, after 2 years, the capacity was 33%, and in the 3rd year, it was 48%. In subsequent years, a decrease in germination was noted, with a 5.5-year period showing a capacity of 6% [24].
As Kukowski [24] proved, diaspores located in the soil may have different germination compared to those stored in the laboratory. Furthermore, the viability was 43–47%, respectively, for diaspores stored in the soil at a depth of 8 cm and 20 cm after 3 months. After 6 months, diaspores stored at an 8 cm depth in soil had a germination rate of 63% and at 20 cm, 75%. After a year of storage, the germination rate was 2–3%, and in the fifth year, it was 6% at both depths [24].
As reported by Holm (1972) [67], fresh seeds are characterized by a certain degree of primary dormancy and therefore germinate at 30–40%. Then, when the seeds are in the soil, conversion from primary to secondary dormancy takes place and the light requirement for germination is acquired. However, in our study, the seeds were not stored in the soil, so the conversion from aerobic to anaerobic metabolism was not possible. They also had no access to light.
The growth habitat of the mother plant is one of the factors cited as affecting seed germination. The fertilization of mother plants is an important factor determining seed quality [68,69]. Both an overabundance of major macronutrients and a deficiency can lead to disorders in seed germination [68].
Nitrogen is an ingredient that stimulates the mother plant to produce sizeable seeds that are high in protein. However, this situation can lead to disorders in germination as a result of problems with delayed maturation. As shown by Sánchez et al. [70], intensive nitrogen fertilization of mother plants may contribute to extending the longevity of the produced seeds, but it may reduce their dormancy. According to the authors’ reports, the germination capacity of seeds produced by the mother plant is varied by the elemental abundance of the soil. As Kristó et al. [71] reported, the germination percentage, seedling health, vigor value, and emergence percentage decreased with the increase in the N dose. However, the authors’ research concerned an agricultural crop (winter wheat). In our study, soil nitrogen content variably influenced the confirmed correlation between soil nitrogen content and diaspore germination, depending on their age. Diaspores of A. spica-venti in the year of harvest and one-year olds showed no significant correlation. The linear relationship was −0.25 and 0.28, respectively. Similar results were recorded by Contreras et al. (2024) [72]. The authors indicate that lack of seed response may be due to low seed dormancy. However, they also noted a significant reduction seed longevity during the cultivation of Chenopodium quinoa Willd, when the highest N fertilization treatment was used.
However, for the older diaspores, a strong correlation was recorded between the germination of A. spica-venti diaspores and the soil N content from the mother plant collection site. The correlations were 0.42, 0.69 and 0.72 for 2-, 3-, and 4-year-old diaspores, respectively. This contradicts the findings of Kapczyńska (2012) [73], where the author showed a decrease in germination of grass seeds with age. One-year-old seeds of such grasses as Melica altissima ‘Atropurpurea’, Melica ciliata ssp. Taurica, Melica transsilvanica, Pennisetum flaccidum exhibited the highest germination percentage. However, with age, the seeds of the studied grass species were characterized by varied vitality. As indicated by the author [73], this may also be caused by species and genetic variability [74]. Unfortunately, the author of [73] provided no information about the growth and development conditions of the mother plant or about the soil’s abundances.
A deficiency of phosphorus in the place where the mother plant grew may lead to lower phosphorus content in its seeds, which may also contribute to their lower germination capacity [69,75,76]. The same is true for potassium [69,77]. This contradicts the opinion of Zhu et al. [78], where the authors indicate that phosphorus deficiency or excessive P application would prolong seed germination (under N0 condition, the mean germination time of seeds from P60 and P120 treatments significantly decreased compared to those from P0 and P90). The authors of [78] proved that the seed germinability of Bromus inermis Leyss. could be maintained at a higher level at the treatment of 100 N kg·ha−1 and 60 P2O5 kg·ha−1, and such doses of nutrients also increased seed longevity by increasing the level of catalase and acid phosphoesterase activities and decreasing the content of malondialdehyde and hydrogen peroxide (H2O2) in aged seeds. In our own research, the effect on the studied grass (diaspores) was variable, depending mainly on the age of the seed. A moderate and strong positive correlation was noted between the percentage of germinated 3- and 4-year-old diaspores and the content of phosphorus and potassium in the soil. The recorded values were, respectively, p < 0.05; r = 0.41; r = 0.44 and r = 0.67; r = 0.95. As Hajcman [57] points out, deficiency of P and K supply to the mother plants, together with a high N supply, can result in the production of P- and K-deficient seeds with a lower capacity for germination.
There are many reports on the positive effects of algae on plants [79,80,81,82,83,84]. Algae most often contribute to an increase in arable crop yield as well as the alleviation of plant stress. The positive impact is due to the effects of phytohormones, amino acids, and lipids [85]. However, there are no reports on the effects of biostimulants based on algae or silicon on weed diaspores. Some studies have focused on the influence of biostimulants based on other ingredients on weed number and weight. According to Sawicka et al. [86], the Asahi SL preparation [sodium para-nitrophenolate, sodium ortho-nitrophenolate, sodium 5-nitroguaiacolate] increases the fresh and dry weight of weeds in potato cultivation in wet years. Moreover, the use of this preparation increased the incidence of Chenopodium album and Sinapiss arvensis. In turn, the combined use of Asahi SL with Insol 7 contributed to the increase in secondary weed infestation with Echinochloa crus-galli and S. arvensis. The increased occurrence of E. crus-galli was explained by improved cation penetration. Similarly, Maziarek et al. [87] proved that the use of biopreparations contributes to an increased weed biomass in spring wheat monoculture. Specifically, the use of Asahi SL resulted in an increase in weed biomass by 28.2% compared to the control and by 19.9% after the application of Nano-Gro [87].
In this study, a significant positive increase in germination was recorded in the year of diaspore collection, but this depended on the place of occurrence of the plant material (Table 3). After one year of storage, a significant decrease in the germination of diaspores was observed where an algae extract (E. maxima) was used. This was the case for the germination of diaspores from three sites out of five. Observation of the 2-year-old diaspore germination showed mainly an increase in germination or germination at the same level as the control diaspores. As indicated by Wawrzyniak et al. [88], the method of storage can be a determinant of germination. However, as indicated Moncada et al. (2011) [89] auxins are one of the major plant growth regulators contained in E. maxima extracts. Therefore, as they proved, seaweed extracts stronger induce seed germination and growth parameters of Vigna radiata. Similar results were recorded by Hernández-Herrera et al. (2023) [90] in observations of soybean germination. The germination percentage of soybeans for all extract concentrations was higher than in the control group. However, the authors themselves point to the small number of studies on the impact of algae extracts on the germination of crops, not to mention weed diaspores.
In this study, when the solution with added silicon was used, the diaspores showed a significant percentage increase in germination in the year of harvesting [the highest increase, depending on the site, was up to 52% of germinated seeds, 23.75 percentage points more, compared to seeds where distilled water was used]. A similar situation was recorded for two-year-old diaspores, where the highest recorded percentage of germinated seeds was 55.5% and the difference in percentage points was 31.
However, reports on the effect of silicon on seed germination are divergent, depending on the plants tested. As reported by Jiang et al. (2023) [91], Si did not have a large effect on increasing the percentage of germinated seeds of Oryza sativa L. However, as the authors showed, this element could increase the sprout and root length of germinated seeds.

5. Conclusions

Currently, most studies have focused on the resistance of A. spica-venti to herbicide active ingredients. However, reports on seed germination are divergent due to a number of factors. In this study, 3-year-old seeds showed the highest percentage of germination. In addition, their germination process was also positively correlated with soil abundance (from sites of mother plant material collection) of macronutrients (N, P, K). The strongest correlations were recorded for 3- and 4-year-old seeds. A. spica-venti, or rather, the seeds of the weed that were soaked in the biostimulants reacted in different ways. As the diaspores gained age, the germination capacity within the harvested seeds varied, depending further on the type of stimulant and the source of the seed. Additionally, seeds from the harvest year treated with the stimulant were characterized by a higher germination percentage. A. spica-venti, is a highly fecund weed, a trait that makes it very common in crops, appearing in large numbers and providing formidable competition to crops, thus causing significant yield losses. Awareness of the vitality of diaspores will allow for better preparation for the regulation and control of the weed in question, in order to prevent yield loss in crops. The theme of diaspore viability is one to be pursued further. Further research should include observations of factors affecting germination, including new stimulants emerging on the agricultural market.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/agronomy15030715/s1, Figures S1–S54: Figure S1–S10. Germination dynamics of A. spica-venti depending on of mother plant occurrence and diaspores age. Figure S11–S15. Correlations of diaspores A. spica-venti germination with N contained in the soil. Figure S16–S20. Correlations of diaspores A. spica-venti germination with P contained in the soil. Figure S21–S25. Correlations of diaspores A. spica-venti germination with K contained in the soil. Figure S26–S30. Correlations of diaspores A. spica-venti germination with macronutrients and pH of soil. Figure S31–S33. Correlations of germination of A. spica-venti diaspores in the harvest year, one year and two years old seeds with the application of an algae preparation with macronutrients contained in the soil. Figure S34–S36. Correlations of germination of A. spica-venti diaspores in the harvest year, one year and two years old seeds with the application of an algae preparation with macronutrients contained in the soil. Figure S37–S39. Correlations of germination of A. spica-venti diaspores in the harvest year, one year and two years old seeds with the application of an algae preparation with macronutrients contained in the soil. Figure S40–S42. Correlations of germination of A. spica-venti diaspores in the harvest year, one year and two years old seeds with the application of an algae preparation with macronutrients contained in the soil. Figure S43–S45. Correlations of germination of A. spica-venti diaspores in the harvest year, one year and two years old seeds with the application of an silicon preparation with macronutrients contained in the soil. Figure S46–S48. Correlations of germination of A. spica-venti diaspores in the harvest year, one year and two years old seeds with the application of an silicon preparation with macronutrients contained in the soil. Figure S49–S51. Correlations of germination of A. spica-venti diaspores in the harvest year, one year and two years old seeds with the application of an silicon preparation with macronutrients contained in the soil. Figure S52–S54. Correlations of germination of A. spica-venti diaspores in the harvest year, one year and two years old seeds with the application of an silicon preparation with macronutrients contained in the soil.

Funding

This work was supported by the Wrocław University of Environmental and Life Sciences (Poland) as the Ph.D. research program Innovative Scientist no. N060/0009/20.

Data Availability Statement

Data will be made available on request.

Conflicts of Interest

The author declares no conflicts of interest.

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Figure 1. Map of Poland with Lower Silesia marked in red (A) in the area where the research was carried out (B) in three study sites (Wrocław, Sucha Wielka, Głuchów Dolny).
Figure 1. Map of Poland with Lower Silesia marked in red (A) in the area where the research was carried out (B) in three study sites (Wrocław, Sucha Wielka, Głuchów Dolny).
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Figure 2. Age-dependent germination of A. spica-venti diaspores. * The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test, and others are.
Figure 2. Age-dependent germination of A. spica-venti diaspores. * The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test, and others are.
Agronomy 15 00715 g002
Table 1. Places of harvest of Apera spica-venti in Lower Silesia, Poland: “x” means, collection of plant material A. spica-venti.
Table 1. Places of harvest of Apera spica-venti in Lower Silesia, Poland: “x” means, collection of plant material A. spica-venti.
Study SitesLocalityGeographical LatitudesYear of Study
201620172019
IWrocław51.11, 17.14x
winter wheat
x
winter wheat
IIWrocław51.11, 17.14 x
winter wheat
x
winter wheat
IIISucha Wielka51.32, 17,16 x
winter wheat
x
winter wheat
IVWrocław51.11, 17.14 x
rye
x
rye
VGłuchów Dolny51.28, 17.12x
winter wheat
x
winter wheat
“x” indicate collection of plant material A. spica-venti.
Table 2. Life test variants **—vegetation chamber [MLR-352H/SANYO].
Table 2. Life test variants **—vegetation chamber [MLR-352H/SANYO].
VariantsDiaspores AgeType of SubstanceNight/DayHumidity
Variant 1diaspores 3 months after harvest
annual and multi-year diaspores
Distilled waterNight 14 h/10 °C
Day 10 h/15 °C
70% humidity
Variant 2diaspores 3 months after harvest
annual and multi-year diaspores
BiostimulantsNight 14 h/10 °C
Day 10 h/15 °C
70% humidity
** Literature [57].
Table 3. Final germination percentage (FGP) of A. spica-venti [%].
Table 3. Final germination percentage (FGP) of A. spica-venti [%].
Lp.Study Sites 120162017201920202021
I 1A 201620.00 BC3 a4-39.75 A b34.50 AB a12.75 C a
VB 20168.75 B a-55.75 A a51.75 A a13.75 B a
 
IIIC 2017- 25.75 B a31.75 A b32.50 A b9.25 B a
IVD 2017-15.25 B a45.00 A a39.50 A ab11.00 B a
IIE 2017-16.00 B a56.25 A a50.25 A a20.50 B a
 
IA 2019--15.25 B b26.50 A a18.50 AB abc
VB 2019--25.00 B a44.00 A a26.50 B ab
IIIC 2019--9.50 A bc12.50 A a10.00 A c
IVD 2019--28.25 A a22.25 A a31.50 A a
IIE 2019--6.00 A c12.00 A a13.50 A bc
1 See Table 1. 2 The research was not carried out in a given year. 3 Statistical analysis made for each site of harvest with years of observation (horizontal analysis, capital letter, and subscript). 4 Statistical analysis made for different sites of harvest in a given year (vertical analysis, superscript with separation by year, seeds of the same age, and small letter). The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test, and others are.
Table 4. The Pearson (r) correlation between the mean levels of N, P, and K in the soil * and pH of soil with diaspores germination.
Table 4. The Pearson (r) correlation between the mean levels of N, P, and K in the soil * and pH of soil with diaspores germination.
Pearson (r)Diaspores
Harvest Year1-Year2-Year3-Year4-Year
SoilN−0.250.28−0.420.690.72
P0.26−0.280.050.410.44
K−0.15−0.230.200.670.95
pH0.17−0.34−0.36−0.57−0.09
* Table A1—the research material (soil) comes from the same sites of the A. spica-venti presented in the article [54].
Table 5. Final germination percentage (FGP) of A. spica-venti diaspores with the use of E. maxima (collection 2019/analysis 2019 and in 2020 and 2021) [%].
Table 5. Final germination percentage (FGP) of A. spica-venti diaspores with the use of E. maxima (collection 2019/analysis 2019 and in 2020 and 2021) [%].
Study Sites201920202021
ControlE. maximaControlE. maximaControlE. maxima
A 201915.25 B1 a220.00 B a26.50 A a13.50 ABC b18.50 ABC b29.00 B a
B 201925.00 A b34.25 A a44.00 A a17.75 AB b26.50 AB b48.75 A a
C 20199.50 BC a11.50 C a12.50 A a8.75 BC a10.00 C b31.50 B a
D 201928.25 A b44.25 A a22.25 A a25.50 A a31.50 A a37.50 AB a
E 20196.00 C b16.50 BC a12.00 A a4.50 C b13.50 BC a7.50 C a
1 up-index—germination analysis within objects with a given formulation. 2 lower index—germination analysis between control and grains with a given preparation in a given year of analysis. The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test, and others are.
Table 6. Final germination percentage (FGP) of A. spica-venti diaspores with the use of silicon (collection 2019/analysis 2019 and in 2020 and 2021) [%].
Table 6. Final germination percentage (FGP) of A. spica-venti diaspores with the use of silicon (collection 2019/analysis 2019 and in 2020 and 2021) [%].
Study Sites201920202021
ControlSilicon ControlSilicon ControlSilicon
A 201915.25 B1 b224.50 B a26.50 A a2.50 CD b18.50 ABC b36.25 B a
B 2019 25.00 A b46.00 A a44.00 A a0.00 D b26.50 AB b54.00 A a
C 20199.50 BC a12.50 C a12.50 A b20.50 B a10.00 C b41.00 B a
D 201928.25 A b52.00 A a22.25 A a37.00 A a31.50 A b55.50 A a
E 20196.00 C b21.75 BC a12.00 A a7.50 C a13.50 BC a11.00 C a
1 up-index—germination analysis within objects with a given formulation. 2 lower index—germination analysis between control and grains with a given preparation in a given year of analysis. The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test; others are.
Table 7. The Pearson (r) correlation between the mean levels of N, P, and K in the soil and pH of soil with A. spica-venti diaspores germination with alge E. maxima.
Table 7. The Pearson (r) correlation between the mean levels of N, P, and K in the soil and pH of soil with A. spica-venti diaspores germination with alge E. maxima.
Pearson (r)Diaspores
201920202021
SoilN−0.45−0.61−0.29
P−0.13−0.18−0.64
K−0.56−0.62−0.70
pH−0.15−0.14−0.15
Table 8. The Pearson (r) correlation between the mean levels of N, P, and K in the soil and pH of soil with A. spica-venti diaspores germination with silicon.
Table 8. The Pearson (r) correlation between the mean levels of N, P, and K in the soil and pH of soil with A. spica-venti diaspores germination with silicon.
Pearson (r)Diaspores
201920202021
SoilN−0.32−0.97−0.55
P−0.15−0.17−0.58
K−0.52−0.62−0.79
pH−0.08−0.71−0.28
Table 9. Length of the first leaf of A. spics-venti [mm]—control.
Table 9. Length of the first leaf of A. spics-venti [mm]—control.
Study Sites201920202021
A 201912.90 A1 a211.88 A ab8.49 AB b
B 201910.87 A a11.39 A a7.25 BC b
C 20199.59 A a10.63 A a6.55 C a
D 201912.09 A a12.40 A a9.18 A a
E 201911.75 A a12.01 A a8.11 AB b
1 up-index—analysis of 1-leaf length within sites in a given year germination. 2 lower index—analysis of 1-leaf length in years within site. The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test; others are.
Table 10. Length of the first leaf of A. spics-venti [mm]—silicon.
Table 10. Length of the first leaf of A. spics-venti [mm]—silicon.
Study Sites201920202021
A 201911.07 B1 a27.15 C c9.59 AB b
B 201910.59 B a0.00 D c9.03 B b
C 20199.08 C a9.47 B a9.33 AB a
D 201912.80 A a9.33 B b10.10 A b
E 201913.56 A a13.49 A a10.02 A b
1 up-index—analysis of 1-leaf length within sites in a given year germination. 2 lower index—analysis of 1-leaf length in years within site. The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test; others are.
Table 11. Length of the first leaf of A. spics-venti [mm]—E. maxima.
Table 11. Length of the first leaf of A. spics-venti [mm]—E. maxima.
Study Sites201920202021
A 201910.91 AB1 a28.75 A b10.69 AB a
B 20199.80 B a6.68 B b9.43 C a
C 20199.66 B a7.87 AB b9.32 C ab
D 201911.84 AB a9.34 A b10.28 B ab
E 201912.78 A a8.11 AB b11.36 A a
1 up-index—analysis of 1-leaf length within sites in a given year germination. 2 lower index—analysis of 1-leaf length in years within site. The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test; others are.
Table 12. Length [mm] of 1-leaf of A. spica-venti from diaspores in 2019, 2020 and 2021 in relation to the effect of the substance (collection 2019/analysis 2019 and in 2020 and 2021).
Table 12. Length [mm] of 1-leaf of A. spica-venti from diaspores in 2019, 2020 and 2021 in relation to the effect of the substance (collection 2019/analysis 2019 and in 2020 and 2021).
Study Sites201920202021
ControlSiliconE. maximaControlSilicon E. maximaControlSilicon E. maxima
A 201912.90 a111.07 a10.91 a11.39 a0.00 c6.68 b8.49 c9.59 b10.64 a
B 201910.87 a10.59 a9.80 a12.40 a9.34 b9.35 b7.25 b9.03 a9.43 a
C 20199.59 a9.08 a9.66 a10.63 a9.47 a7.87 b6.55 b9.32 a9.38 a
D 201912.10 a12.80 a11.43 a11.88 a7.15 c8.75 b9.18 b10.10 a10.28 a
E 201911.75 b13.56 a12.78 ab12.01 a13.49 a8.11 b8.10 b10.03 a11.36 a
1 up-index—analysis of 1-leaf length within sites in a given year germination. The same letter means that results are not statistically different at the α ≤ 0.05 level, according to Tukey’s HSD test; others are.
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Lejman, A. The Relationship Between the Germination of Silky Bent Grass (Apera spica-venti (L.) Beauv.) Diaspores and Their Age, Place of Occurrence, and Action of Stimulating Substances. Agronomy 2025, 15, 715. https://doi.org/10.3390/agronomy15030715

AMA Style

Lejman A. The Relationship Between the Germination of Silky Bent Grass (Apera spica-venti (L.) Beauv.) Diaspores and Their Age, Place of Occurrence, and Action of Stimulating Substances. Agronomy. 2025; 15(3):715. https://doi.org/10.3390/agronomy15030715

Chicago/Turabian Style

Lejman, Agnieszka. 2025. "The Relationship Between the Germination of Silky Bent Grass (Apera spica-venti (L.) Beauv.) Diaspores and Their Age, Place of Occurrence, and Action of Stimulating Substances" Agronomy 15, no. 3: 715. https://doi.org/10.3390/agronomy15030715

APA Style

Lejman, A. (2025). The Relationship Between the Germination of Silky Bent Grass (Apera spica-venti (L.) Beauv.) Diaspores and Their Age, Place of Occurrence, and Action of Stimulating Substances. Agronomy, 15(3), 715. https://doi.org/10.3390/agronomy15030715

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