Skip to main content
NCBI home page
BMC Plant Biology logoLink to BMC Plant Biology
. 2025 Sep 25;25:1222. doi: 10.1186/s12870-025-07244-9

Effects of whole-plant extracts of four species dominant in the Qinghai-Tibetan plateau on their germination and growth patterns

Hui Li 1, Jianpeng Wang 1, Yumao Ning 1, Shree Prakash Pandey 1, Zhenqing Li 2, Yongjie Liu 1,✉
PMCID: PMC12465637  PMID: 40999332

Abstract

Allelopathy plays an important role in driving plant interactions. Effects of allelopathy on seed germination and plant growth have been extensively investigated, while the impacts of whole-plant water extracts of different plants on their seed germination, and growth of shoots and roots are still unclear. Therefore, a manipulation experiment was conducted to explore such effects of whole-plant extract of four dominant species of the Qinghai-Tibetan Plateau (Elymus nutans, Festuca sinensis, Poa pratensis and Vicia unijuga). Four extract concentrations (0, 0.025, 0.05, and 0.1 g/ml) of each species were applied on the seeds of the four species. Results showed that water extract significantly affected their seed germination rate, and shoot length and root length. Specifically, the seed germination rate and seed germination index decreased with increasing extract concentrations, while the mean germination time increased. Further, the medium time of germination (the peak value of the germination period) had postponed with increasing extract concentrations. Moreover, the root length decreased with increasing extract concentration, yet the shoot length was only inhibited by the highest extract concentration. A stronger effect of the extracts on the roots than on the shoots was also inferred since (1) extract concentration was negatively correlated with the root length (not shoot length), and (2) root length contributed more than shoot length to the synthetical allelopathic effect index. Thus, root length could be a better bio-indicator in allelopathy assays than shoot length. Also, the plant species composition should be considered for obtaining significant effects of allelochemicals on seed germination and plant growth.

Keywords: Allelopathy, Concentration, Shoot length, Root length, Seed germination, Water extract

Introduction

Allelopathy is a process where plants release allelochemicals that affect the establishment and growth of their neighbours [1, 2]. Plants normally release allelochemicals into the environment through four pathways, leaching through rain, plant litter decomposition, roots exudation, and leaf volatilization [3]. Allelochemicals, such as ferulic acid, usually affect seed germination and plant growth [4–7]. Allelochemicals could be released from plants into the environment with the help of leaching after rains [3]. However, allelochemicals that are released by leaching due to rain in natural condition are difficult to detect due to the complex nature of the leached mixture. Thus, many studies investigate them by applying water extract of the shoots, roots or the whole plant so that such a process could be stimulated [8–10].

Water extract is widely used to study allelopathy [11–13]. Some studies found that higher extract concentration of Rhododendron capitatum litters significantly repressed seed germination of its neighbors (Elymus nutans, Poa pratensis, and Medicago ruthenica) [14]. Similarly, some other studies indicated that the seed germination of Lactuca sativa was inhibited by various concentrations of the leaf extracts of Solidago canadensis [15], and the growth of Amygdalus pedunculata were inhibited by various concentrations of the leaf extracts of Pinus sylvestris [16]. The seed germination of Leymus chinensis had decreased with an application of an increasing shoot extract concentration of E. nutans [17]. Plant height and root length of Tagetes erecta decreased with the application of increasing root extract concentration of Rhus typhina [18]. However, some other studies have revealed that water extracts of Amorpha fruticosa, Hedysarum mongolicum and Sabina vulgaris showed a pattern of promotion of the plant growth at low concentration, whereas their high concentrations exhibited inhibition on Amygdalus pedunculata [19]. In addition, inhibition of conspecies neighbours as an example of allelopathy is also reported [20]. Water extracts of plants not only affect their neighbours (e.g. seed germination and plant growth) but also on themselves [1, 21, 22]. Application of water extracts could promote the generation of reactive oxygen species (ROS) and decrease the ROS scavenging capacity [23]. This could cause oxidative damage, which inhibits the seed germination and plant growth [24]. How allelopathy shape the plant community structure in Qinghai-Tibetan Plateau, which is an important part in the global terrestrial ecosystem [25], remain active research [26].

Elymus nutans, Festuca sinensis, Poa pratensis and Vicia unijuga are the four dominant plant species in the Qinghai-Tibetan Plateau. However, there are relatively few reports that have investigated the effect of varying whole-plant water extract concentrations on the seed germination, shoot length and root length of these dominant plant species. Accordingly, a manipulation experiment was conducted to explore the effects of various concentrations of whole-plant extracts of the four plant species (E. nutans, F. sinensis, P. pratensis and V. unijuga) on their seed germination and plant growth. Four concentrations (0, 0.025, 0.05 and 0.1 g/mL) of each of the four extracts were developed. The following specific assumptions are made: (1) seed germination is assumed to decrease with the increasing water extract concentration since these extracts could inhibit the water uptake of seeds and then reduce the seed germination [27]. (2) Root length is assumed to decrease with increasing extract concentration as the allelochemicals in the water extract could inhibit cell proliferation in the root apical meristem [28] and the higher extract concentration could have more negative effect than lower water extract concentration on root length [16]. (3) Shoot length is expected to be shortened by the application of the relatively higher extract concentration as the extract could constrain the activity of antioxidant enzymes [29], reducing the scavenging effect on ROS and destroying the membrane system of plants to inhibit the shoot growth [30–32]. In general, the abovementioned effects could be modified by the origins of water extracts [33, 34].

Materials and methods

To explore the effects of whole-plant extract concentrations of the four dominant plant species on the seed germination, shoot length and root length of these plant species, an experiment was conducted in a greenhouse at the Yuzhong Campus of Lanzhou University, China (104°09′44″E, 35°56′55″N) from November 2023 to March 2024. This study included three steps, i.e. growing the four plant species, preparing water extracts of these plants, and exploring effects of these water extracts on the seed germination and plant growth.

Step 1 Four plant species that are dominant in the Qinghai-Tibetan Plateau (E. nutans, F. sinensis, P. pratensis, V. unijuga, labelled as EN, FS, PP and VU, respectively) were planted in pots with disinfected sand in the greenhouse on 8 November, 2023. All the seeds in this study were collected by Jianquan Zhang from the natural grasslands that located in Luqu County in the Gansu Province in the Qinghai-Tibetan Plateau.

Step 2 After 60 days (on 7 January 2024) of planting, the whole plant from each pot was harvested and stored in a cool and dry place to air-dry. The air-dried plants were crushed and set aside for preparation of water extracts. Specifically, 20 g of the air-dried plants of the four species were powdered and placed in 200 mL of pure water in a conical flask. Then, they were shaken for 2 h every 12 h at a speed of 200 revolutions per minute. After 48 h, negative air pressure was used to filter the mixtures to obtain the initial water extracts [8]. These initial extracts were diluted to 0.05 g/mL and 0.025 g/mL with pure water. Finally, the water extracts of the four plant species were stored in brown bottles at 4 °C until use.

Step 3 To assess effects of these water extracts on the seed germination, shoot length and root length of these four plant species, 4 forage plant seeds and 12 treatments (water extracts of 4 species, E. nutans, F. sinensis, P. pratensis and V. unijuga, each of the concentrations of 0, 0.025, 0.05 and 0.1 g/mL) were applied in this step, and pure water (0 g/mL) was used as the control (Fig. 1), each treatment had 3 replicates. To break the dormancy, the seeds of V. unijuga were treated with concentrated (98% v/v) sulfuric acid for 15 min [35]. Seeds of the four plant species were surface sterilized with 1% (v/v) sodium hypochlorite for 15 min and rinsed with pure water [36]. Then, 30 healthy sterile seeds of each plant species were placed in petri dishes with a diameter of 9 cm containing two layers of filter papers soaked with either 5 mL water extract or pure water [8]. The dishes were then sealed with parafilm to reduce water loss and set in an incubator chamber (25 °C; 12 h/12 h light/dark cycle; 4000 lx). The following measurements were conducted in this step:

Fig. 1.

Fig. 1

Open in a new tab

Setup of the experiment in this study, and it includes three stpes, where step 1 refers to growing the four plant species in a greenhouse (a), step 2 refers to using these plants generating the four extrat concentrations (b), and step 3 refers to treating these extract concentrations on the seeds of these four plant species (c), where these treatments were put into incubator chambers

Germinated seeds were enumerated on a daily basis. Seeds were scored as germinated if the length of their radicle was 1–2 mm [37]. After 3 weeks, 5 plants per petri dish were randomly selected to measure their shoot length and root length. After measuring the shoot length and root length, the five selected plants were pool together in an alluminium foil, and immediately frozen in liquid nitrogen, and stored in −80 °C until further use. This constituted one biological replicate, and 5 such replicates were collected. 0.1 –0.2 g of each replicate was used to measure the activities of superoxide dismutase (SOD), catalase (CAT), peroxidase (POD) and the content of soluble protein (SP).

The following plant indices were quantified:

(1) Germination indices.

graphic file with name d33e519.gif
graphic file with name d33e525.gif
graphic file with name d33e531.gif
graphic file with name d33e537.gif

Where 30 is the total number of seeds in the petri dishes, Gt is the number of germinated seeds at time T, and Dt is the number of the days after planting [8, 38, 39].

(2) Growth indices.

To measure the shoot length and root length, five seedlings were randomly selected in each dish, and the average values of these selected seedlings in each dish were applied in the following analyses. To explore the effect of water extracts on the physiological responses of the seedlings, activities of superoxide dismutase (SOD), catalase (CAT), and peroxidase (POD) and soluble protein (SP) contents were measured. Specifically, the SOD activity was measured with a colorimetric assay by using Nitro blue tetrazolium illumination [40, 41], and the POD activity was measured with the help of a colorimetric assay using guaiacol [42, 43], whereas CAT activity was measured with the help of an ultraviolet spectrophotometry [44]. Total SP content was measured with a colorimetric assay using Coomassie brilliant blue G-250 staining [45, 46].

(3) Allelopathy indices.

graphic file with name d33e601.gif
graphic file with name d33e609.gif

Where C and T are the control and treatment values, respectively, GR, GI and MGT are the seed germination rate, seed germination index and mean germination time, respectively. SL and RL are the shoot length and root length, respectively [16, 47].

Statistical analyses

To investigate the effects of water extracts of the four plant species on their seed germination, shoot length and root length, analyses were conducted at two scales, i.e. at a combination scale, where the four types of water extracts were combined, and at species scale, where the water extracts of the four plant species were separated.

At the combination scale, (1) a general linear model (GLM) was used to explore the effects of water extracts, extract concentrations, and their interaction on the seed germination rate, shoot length and root length. (2) Person’s correlation analysis was used to test the relationships between the extract concentration and the seed germination, shoot length, root length, SOD, CAT, POD and SP. Note that SOD, CAT, POD and SP were derived from the whole plants including both shoots and roots of E. nutans, F. sinensis and P. pratensis. (3) A random forest analysis was done to explore the contribution of seed germination (seed germination rate, seed germination index and mean germination time), shoot length, root length, SOD, CAT, POD and SP of the four plant species to the synthetical allelopathic effect index (SE).

At the species scale, a GLM was used to explore the effects of the extract concentrations of the four plant species on their seed germination rate, shoot length and root length.

Results

Effects of whole-plant extracts on the seed germination

At the combination scale, seed germinations of E. nutans, F. sinensis, P. pratensis and V. unijuga were significantly affected by the various extract concentrations (Table 1). The seed germination rates of the four plant species decreased with increasing extract concentrations (Fig. 2a). Further, the seed germination index also decreased with the increasing extract concentrations (Fig. 2b), and the mean germination time had increased (Fig. 2c). Besides, the daily seed germination rate had also decreased along the germination date. Moreover, the peak value of germination period had decreased (compared that under extract concentration of 0.025 g/mL and 0.1 g/mL, it decreased by 92%), and the peak value of germination period was postponed (0.025 g/mL vs. 0.1 g/mL, it postponed by 7 days, Fig. 2d).

Table 1.

At the combination scale, effects of water extracts of the four plant species (Elymus nutans, Festuca sinensis, Poa pratensis and Vicia unijuga) dominant in the Qinghai-Tibetan plateau, extract concentrations (0, 0.025, 0.05 and 0.01 g/mL), and their interaction in general linear model (GLM) on the seed germination rate, shoot length and root length of the four plant species.

Source df F P
Seed germination rate
 Water extract 3 9.72 < 0.001
 Extract concentration 2 42.793 < 0.001
 Water extract × Extract concentration 6 2.067 0.061
Shoot length
 Water extract 3 3.904 0.01
 Extract concentration 2 23.914 < 0.001
 Water extract × Extract concentration 6 1.874 0.089
Root length
 Water extract 3 9.069 < 0.001
 Extract concentration 2 24.684 < 0.001
 Water extract × Extract concentration 6 1.625 0.144
Open in a new tab

F-values, P-values and degrees of freedom (df) are given, with significant results (P < 0.05) in bold

Fig. 2.

Fig. 2

Open in a new tab

At the combination scale, effects of extract concentrations of the four plant species (Elymus nutans, Festuca sinensis, Poa pratensis and Vicia unijuga, labelled as EN, FS, PP and VU, respectively) on their seed germination rate (a), seed germination index (b), mean germination time (c) and daily seed germination rate (d). Different letters indicate differences among the extract concentrations. The red lines in (d) indicate the peak value of germination period

At the species scale, the extracts of different species had similar effects on the seed germination rates of E. nutans, F. sinensis, P. pratensis and V. unijuga (Tables 2, 3, 4 and 5). The seed germination rates of the four plant species decreased with increasing extract concentrations of different plant species (Fig. 3).

Table 2.

At the species scale, effects of extract concentrations (0, 0.025, 0.05 and 0.01 g/mL) of Elymus nutans in a general linear model (GLM) on the seed germination rate, shoot length and root length of the four target plant species (Elymus nutans, Festuca sinensis, Poa pratensis and Vicia unijuga)

Source df F P
Seed germination rate
 Extract concentration 2 69.324 < 0.001
 Species 3 50.525 < 0.001
 Extract concentration × Species 6 15.32 < 0.001
Shoot length
 Extract concentration 2 7.697 0.003
 Species 3 9.806 < 0.001
 Extract concentration × Species 6 1.227 0.329
Root length
 Extract concentration 2 9.751 0.001
 Species 3 6.66 0.002
 Extract concentration × Species 6 2.471 0.054
Open in a new tab

F-values, P-values and degrees of freedom (df) are given, with significant results (P < 0.05) in bold

Table 3.

At the species scale, effects of extract concentrations (0, 0.025, 0.05 and 0.01 g/mL) of Festuca sinensis in a general linear model (GLM) on the seed germination rate, shoot length and root length of the four target plant species (Elymus nutans, Festuca sinensis, Poa pratensis and Vicia unijuga)

Source df F P
Seed germination rate
 Extract concentration 2 45.741 < 0.001
 Species 3 17.21 < 0.001
 Extract concentration × Species 6 4.593 < 0.001
Shoot length
 Extract concentration 2 24.546 < 0.001
 Species 3 40.389 < 0.001
 Extract concentration × Species 6 4.653 0.003
Root length
 Extract concentration 2 42.61 < 0.001
 Species 3 23.366 < 0.001
 Extract concentration × Species 6 3.244 0.018
Open in a new tab

F-values, P-values and degrees of freedom (df) are given, with significant results (P < 0.05) in bold

Table 4.

At the species scale, effects of extract concentrations (0, 0.025, 0.05 and 0.01 g/mL) of Poa pratensis in a general linear model (GLM) on the seed germination rate, shoot length and root length of the four target plant species (Elymus nutans, Festuca sinensis, Poa pratensis and Vicia unijuga)

Source df F P
Seed germination rate
 Extract concentration 2 90.195 < 0.001
 Species 3 44.227 < 0.001
 Extract concentration × Species 6 16.002 < 0.001
Shoot length
 Extract concentration 2 97.014 < 0.001
 Species 3 48.788 < 0.001
 Extract concentration × Species 6 12.856 < 0.001
Root length
 Extract concentration 2 72.052 < 0.001
 Species 3 27.951 < 0.001
 Extract concentration × Species 6 8.436 < 0.001
Open in a new tab

F-values, P-values and degrees of freedom (df) are given, with significant results (P < 0.05) in bold

Table 5.

At the species scale, effects of extract concentrations (0, 0.025, 0.05 and 0.01 g/mL) of Vicia Unijuga in a general linear model (GLM) on the seed germination rate, shoot length and root length of the four target plant species (Elymus nutans, Festuca sinensis, Poa pratensis and Vicia Unijuga)

Source df F P
Seed germination rate
 Extract concentration 2 40.692 < 0.001
 Species 3 6.504 0.002
 Extract concentration × Species 6 6.504 < 0.001
Shoot length
 Extract concentration 2 32.57 < 0.001
 Species 3 8.927 < 0.001
 Extract concentration × Species 6 8.927 < 0.001
Root length
 Extract concentration 2 3.851 0.035
 Species 3 7.416 0.001
 Extract concentration × Species 6 2.157 0.084
Open in a new tab

F-values, P-values and degrees of freedom (df) are given, with significant results (P < 0.05) in bold

Fig. 3.

Fig. 3

Open in a new tab

At the species scale, effects of various concentrations of the water extracts on the seed germination rates. Left panel is the pictural representation of design of the experiment, and right panel is the quantitative evaluations. Evaluation of 4 extract concentrations of Elymus nutans (a), Festuca sinensis (b), Poa pratensis (c) and Vicia unijuga (d) are tested on all the 4 species (labelled as EN, FS, PP, and VU, respectively). Different letters indicate significance of differences of 4 extract concentrations in a species

Note that the mortalities of E. nutans, F. sinensis, P. pratensis and V. unijuga in the controls were 35%, 3%, 37% and 29%, respectively.

Effects of whole-plant extracts on the shoot length

At the combination scale, shoot lengths of E. nutans, F. sinensis and P. pratensis were significantly inhibited at the highest extract concentration (Table 1; Fig. 4a).

Fig. 4.

Fig. 4

Open in a new tab

At the combination scale, effects of extract concentration on the shoot length (a) and root length (b) of E. nutans, F. sinensis and P. pratensis, where the germination rate of V. unijuga was too low to quantify its shoot length and root length. Note that Elymus nutans, Festuca sinensis and Poa pratensis are labelled as EN, FS and PP, respectively. Different letters indicate differences between the shoot length or root length of plants treated by different extract concentrations

At the species scale, the extracts of the four plant species had different effects on the shoot length of E. nutans, F. sinensis and P. pratensis (Tables 2, 3, 4 and 5). Specifically, the shoot lengths of E. nutans and F. sinensis were significantly inhibited by the highest water extract concentrations of the four plant species, while the shoot length of P. pratensis was not significantly affected by the extract of F. sinensis (Fig. 5). The shoot length of E. nutans was not significantly affected by the 0.025 g/mL extract of P. pratensis, while the shoot length of F. sinensis and P. pratensis were significantly improved by the 0.025 g/mL extract of P. pratensis (Fig. 5c).

Fig. 5.

Fig. 5

Open in a new tab

At the species scale, effects of extract concentrations of Elymus nutans (a, b), Festuca sinensis (c, d), Poa pratensis (e, f) and Vicia unijuga (g, h) on the shoot length and root length of the E. nutans, F. sinensis and P. pratensis. Note that Elymus nutans, Festuca sinensis and Poa pratensis are labelled as EN, FS and PP, respectively. Different letters indicate differences between the shoot length or root length of plants treated by extract concentrations of E. nutans, F. sinensis, P. pratensis and Vicia unijuga, where the germination rate of V. unijuga was too low to quantify its shoot length and root length

Effects of whole-plant extracts on the root length

At the combination scale, root lengths of the three plant species were significantly decreased by the application of the extracts (Table 1; Fig. 4b).

At the species scale, the extracts of the four plant species had similar effects on the root length of E. nutans, F. sinensis and P. pratensis (Tables 2, 3, 4 and 5), and they were inhibited with increasing extract concentrations (Fig. 5).

Specifically, at the combination scale, the root length was significantly negatively correlated with water extract concentration, the SP content, and the activities of POD and SOD (Fig. 6). The shoot length was however, not negatively correlated with the use of the extract concentrations (Fig. 6). Significant differences of SE were found among the extract concentrations of the four plant species, where the highest extract concentration had stronger inhibited effects on the seed germination and plant growth than the rest extract concentrations, and such significance was found among all the four plant species (Fig. 7). Moreover, the random forest analysis of the importance of the characteristics on the SE demonstrated that root length was more important than the shoot length, and the activities of SOD and POD (Fig. 8). Furthermore, the random forest analyses of the importance of the characteristics on the SE demonstrated that seed germination index was the most important characteristic, and the root length was more important than shoot length, content of SP, activity of SOD, POD and CAT (Fig. 8).

Fig. 6.

Fig. 6

Open in a new tab

At the combiantion scale, person’s correlation analysis of the seed germination rate, seed germination index, mean germination time, shoot length, root length, activities of superoxide dismutase, catalase and peroxidase, content of soluble protein, synthetical allelopathic effect index and water extract concentrations (labelled as GR, GI, MGT, SL, RL, SOD, CAT, POD, SP, SE and C, respectively), where the blue and red represent a negative and positive relationship, respectively, and the darker the color, the stronger the correlation

Fig. 7.

Fig. 7

Open in a new tab

At the combination scale, synthetical allelopathic effects of extract concentrations of the four plant species (Elymus nutans, Festuca sinensis, Poa pratensis and Vicia unijuga, labelled as EN, FS, PP and VU, respectively) on seed germination rate, seed germination index, mean germination time, shoot length, root length, activities of superoxide dismutase, catalase and peroxidase, content of soluble protein. In this case, the synthetical allelopathic effect index (SE) value was calculated as the average allelopathy response index (RI) value of all indicators (seed germination rate, seed germination index, mean germination time, shoot length, root length, activities of superoxide dismutase, catalase and peroxidase, content of soluble protein) of the four plant species treated by the extract concentrations of the four plant species. Different letters indicate differences between the SE of plants treated by different extract concentrations

Fig. 8.

Fig. 8

Open in a new tab

At the combination scale, the importance of seed germination rate, seed germination index, mean germination time, shoot length, root length, activities of superoxide dismutase, catalase and peroxidase and content of soluble protein (marked as GR, GI, MGT, SL, RL, SOD, CAT, POD and SP, respectively) on the synthetical allelopathic effect index (SE) by random forest analysis. The importance from the most to the least of these parameters is listed from top to bottom

Discussion

Water extract significantly affected the seed germination, shoot length and root length. Specifically, seed germination rate and root length decreased with increasing extract concentration, while shoot length was significantly inhibited only at the highest extract concentrations.

According to our first hypothesis, seed germination was assumed to decrease with increasing extract concentrations. This hypothesis was supported as we found that seed germination rates of the four plant species decreased with increasing extract concentration. Such result could be explained by that the higher extract concentration had inhibited seed germination mainly through inhibiting radicle growth and disturbing the balance between the production and scavenging of ROS [24, 48, 49]. Moreover, in the current experiment, the mean germination time increased, while the peak value of germination period was postponed with increasing extract concentration. Such results could be explained by the observation that the higher water extract concentration inhibited water uptake of seeds during the germination period, which had pronged the period of seed imbibition [27, 50, 51]. As a result, a delay in the beginning of seed germination was observed.

According to our second hypothesis, root length is assumed to decrease with the increasing water extract concentration. This hypothesis was supported as we found that the root length of the three target species had decreased with an increase in the extract concentrations. Previous studies have confirmed that allelochemicals, such as volatile monoterpenes, eucalyptol and camphor, had inhibited the root length by widening and shortening the root cells [52, 53], or inhibiting cell proliferation and DNA synthesis in the root apical meristem [28, 54]. The water extract concentrations of 0.025, 0.05 and 0.1 g/mL had significantly inhibited the root length, while only the highest water extract concentration (0.1 g/mL) could significantly inhibit the shoot length. The extract concentration was strongly negatively correlated with the root length, but not significantly negatively correlated with shoot length. Those observations demonstrate that plant roots are more sensitive to the water extract concentration. Previous studies have indicated that water extract had different allelopathic effects on different organs of the plants [14, 29]. The higher sensitivity of root growth is probably because root is the first plant organ to absorb allelochemicals from water extracts, and root cell is more permeable by its nature than the cells of the other plant organ [28, 55, 56].

According to our third hypothesis, the shoot length is expected to be impeded by higher water extract concentration. This hypothesis was partly supported as we found that the shoot length is inhibited by the highest water extract concentration, while such pattern was not found in P. pratensis. Previous studies found that water extracts inhibited shoot length via inhibiting the activity of antioxidant enzymes [29], reducing the scavenging effect on ROS, and destroying the whole membrane system of the plants, as a result it inhibited the plant growth [30, 31, 57]. Meanwhile, the negative effect of water extract on shoot length is related with the negative effect on plant root, where water extract might affect the uptake of water and nutrients by roots [58], then indirectly influence shoot length. However, only the highest concentration inhibited shoot length of E. nutans and F. sinensis in the current study. Such difference could be explained by that the effect of water extract depends on the concentration, and significant differences of SE supported the highest extract concentration had stronger inhibited effects on the plant growth than the rest extract concentrations. The shoot length can inhibit when the water extract reaches a higher concentration, this is consistent with previous research [14, 18]. Moreover, the shoot length of P. pratensis was not impacted by the water extract concentration of F. sinensis, and such result could be derived from (1) the low water extract concentration, where 0.1 g/mL might not be high enough to limit the shoot length of P. pratensis; or (2) the specific-species trait, where studies found that some plant species could resist the negative effects of allelopathy on plant growth [59, 60]; or (3) adding water extracts may increase plant nutrient availability, which partly offsets the inhibition of plant growth by allelochemicals [3].

We founded the activity of SOD and SP contents had increased with increasing extract concentration (Appendix Fig. 1, one-way ANOVA, P < 0.05). The activity of POD and CAT were not impacted by the extract concentration (Appendix Fig. 1, one-way ANOVA, P < 0.05). Such result might be derived from the following reasons: (1) the short-period experiment in this study might not be long enough to detect the effects of extract concentrations on the activity of POD and CAT enzymes since POD and CAT would be activated by enough H2O2, which was conversed by ROS with the help of SOD, and both of these processes take time and may not work within a short period [61–63]. (2) H2O2 could be decomposed by enzymes such as ascorbate peroxidase, leading POD and CAT might not be activated by enough H2O2 [64]. The extract concentration was strongly positively correlated with the activity of SOD (Fig. 6). After the exposure to water extracts and allelochemicals, plants may rapidly produce ROS in the contact area [31, 65]. Then, the first line of defense against the ROS had induced the damage and the activity of SOD increased in the plants that were treated with water extract [29]. However, SOD cannot remove all ROS, and the accumulated ROS could still destroy the plant cells, and thus would have inhibited the plant growth.

The results of this work should be interpreted and extrapolated with caution due to the following reasons: (1) only three extract concentrations were applied in this experiment, and more extract concentrations should be considered to better understand the responses of seed germination and plant growth to allelopathy [66]. (2) Plant endogenous hormones regulate seed germination and plant growth [67, 68]. Allelochemicals can destroy the plant protective enzyme system and the balance of plant endogenous hormones [69]. Therefore, plant endogenous hormones should be considered when exploring the influence of allelopathy. (3) Water extract was generated from the whole plant including both shoot and root in this study, while allelopathy may have different effects on physiological indicators of plant shoots and roots [28, 70]. Thus, physiological indicators of shoots and roots should be separately measured in the future.

Conclusion

Water extract significantly inhibited seed germination and plant growth. Specifically, seed germination rate decreased with increasing water extract concentration, and peak value of germination period was postponed with increasing extract concentration. Root length decreased with increasing extract concentration, while shoot length inhibited by the highest extract concentration. Plant root is more sensitive than shoot to the inhibition by the extract concentration. Therefore, root length could be a better bio-indicator in allelopathy assays than shoot length, and plant species composition should be considered in terms of the significant effects of allelochemicals on seed germination and plant growth.

Acknowledgements

We acknowledge the assistance of Yan Zhang and Mingrui Liu during the experiment.

Abbreviations

ROS

Reactive oxygen species

SOD

Superoxide dismutase

CAT

Catalase

POD

Peroxidase

SP

Soluble protein

GR

Seed germination rate

GI

Seed germination index

MGT

Mean germination time

SE

Synthetical allelopathic effect index

Appendix

See Fig. 9.

Fig. 9.

Fig. 9

Open in a new tab

At the combinatioin scale, effects of extract concentration of the three target plant species (E. nutans, F. sinensis and P. pratensis) on the activities of superoxide dismutase (SOD, a), peroxidase (POD, b), catalase (CAT, c) and content of soluble protein (SP, d). Different small letters indicate differences of plants treated by extract concentrations of E. nutans, F. sinensis, P. pratensis and V. unijuga, where the germination rate of V. unijuga was too low to quantify its activities of SOD, POD, CAT and content of soluble protein

Author contributions

YL designed the study. HL, JW, YN and YL conducted the experiment, collected the data and did the analyses. HL, YL, ZL and SP wrote the draft of the manuscript. All authors contribute substantially to this work.

Funding

This work was supported by the Fundamental Research Funds for the Central Universities (lzujbky-2025-14).

Data availability

The dataset involved in this study are available with the following link: https://data.4tu.nl/private_datasets/3OjYjT4ODjjtxdnkXw2aGJr8fqUEBrkVsfiiThs93Pk.

Declarations

Ethics approval and consent to participate

Not applicable. Seeds of E. nutans, F. sinensis, P. pratensis, and V. unijuga were collected from the grasslands in Gansu Province in the Qinghai-Tibetan Plateau and were identified by Jianquan Zhang from Lanzhou University, and the collection was permitted by the local government. The voucher specimens for these plants are reserved in Lanzhou University, with voucher ID LZUQTCSPJC, LZUQTZHYM, LZUQTCDZSH, LZUQTWTC, and are available on request.

Consent for publication

Not applicable.

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

References

  • 1.Friedman J, Waller GR. Allelopathy and autotoxicity. Trends Biochem Sci. 1985;10(2):47–50. 10.1016/0968-0004(85)90224-5. [Google Scholar]
  • 2.Meiners S, Kong C, Ladwig L, Pisula N, Lang K. Developing an ecological context for allelopathy. Plant Ecol. 2012;213:1221–7. 10.1007/s11258-012-0078-5. [Google Scholar]
  • 3.Zhang Z, Liu Y, Yuan L, Weber E, van Kleunen M. Effect of allelopathy on plant performance: a meta-analysis. Ecol Lett. 2021;24(2):348–62. 10.1111/ele.13627. [DOI] [PubMed] [Google Scholar]
  • 4.Williams RD, Hoagland RE. The effects of naturally occurring phenolic compounds on seed germination. Weed Sci. 1982;30(2):206–12. 10.1017/S0043174500062342. [Google Scholar]
  • 5.Rasmussen JA, Einhellig FA. Synergistic inhibitory effects of p-coumaric and ferulic acids on germination and growth of grain sorghum. J Chem Ecol. 1977;3:197–205. 10.1007/BF00994146. [Google Scholar]
  • 6.Blum U, Dalton BR, Rawlings JO. Effects of ferulic acid and some of its microbial metabolic products on radicle growth of cucumber. J Chem Ecol. 1984;10:1169–91. 10.1007/BF00988547. [DOI] [PubMed] [Google Scholar]
  • 7.Li Z, Wang Q, Ruan X, Pan C, Jiang D. Phenolics and plant allelopathy. Molecules. 2010;15(12):8933–52. 10.3390/molecules15128933. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 8.Wang K, Wang T, Ren C, Dou P, Miao Z, Liu X, Huang D, Wang K. Aqueous extracts of three herbs allelopathically inhibit lettuce germination but promote seedling growth at low concentrations. Plants. 2022;11(4):486. 10.3390/plants11040486. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 9.Yang S, Zheng Y, Guo Y, Cen Z, Dong Y. Allelopathic effect of phenolic acids in various extracts of wheat against fusarium wilt in Faba bean. Funct Plant Biol. 2023;50(12):1062–72. 10.1071/FP23052. [DOI] [PubMed] [Google Scholar]
  • 10.Gruľová D, Baranová B, Eliašová A, Brun C, Fejér J, Kron I, Campone L, Pagliari S, Nastišin Ľ, Sedlák V. Does the invasive Heracleum mantegazzianum influence other species by allelopathy? Plants. 2024;13(10):1333. 10.3390/plants13101333. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 11.Basile A, Sorbo S, Giordano S, Ricciardi L, Ferrara S, Montesano D, Castaldo Cobianchi R, Vuotto ML, Ferrara L. Antibacterial and allelopathic activity of extract from Castanea sativa leaves. Fitoterapia. 2000;71:S110–6. 10.1016/S0367-326X(00)00185-4. [DOI] [PubMed] [Google Scholar]
  • 12.Li J, Zhao T, Chen L, Chen H, Luo D, Chen C, Miao Y, Liu D. Artemisia argyi allelopathy: a generalist compromises hormone balance, element absorption, and photosynthesis of receptor plants. BMC Plant Biol. 2022;22(1):368. 10.1186/s12870-022-03757-9. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 13.Patanè C, Pellegrino A, Cosentino SL, Testa G. Allelopathic effects of Cannabis sativa L. aqueous leaf extracts on seed germination and seedling growth in durum wheat and barley. Agronomy. 2023;13(2):454. 10.3390/agronomy13020454. [Google Scholar]
  • 14.Yang H, Zhao Y, Wei S, Yu X. Isolation of allelochemicals from Rhododendron capitatum and their allelopathy on three perennial herbaceous plants. Plants. 2024;13(18):2585. 10.3390/plants13182585. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 15.Wu R, Wu B, Cheng H, Wang S, Wei M, Wang C. Drought enhanced the allelopathy of goldenrod on the seed germination and seedling growth performance of lettuce. Pol J Environ Stud. 2020;30(1):423–32. 10.15244/PJOES/122691. [Google Scholar]
  • 16.Wang X, Zhang R, Wang J, Di L, Wang H, Sikdar A. The effects of leaf extracts of four tree species on Amygdalus pedunculata seedlings growth. Front Plant Sci. 2021;11:587579. 10.3389/fpls.2020.587579. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 17.Wang K, Dou P, Miao Z, Huang J, Gao Q, Guo L, Liu K, Rong Y, Huang D, Wang K. Seed germination and seedling growth response of Leymus chinensis to the allelopathic influence of grassland plants. Oecologia. 2024;204:899–913. 10.1007/s00442-024-05539-6. [DOI] [PubMed] [Google Scholar]
  • 18.Qu T, Du X, Peng Y, Guo W, Zhao C, Losapio G. Invasive species allelopathy decreases plant growth and soil microbial activity. PLoS ONE. 2021;16(2):e0246685. 10.1371/journal.pone.0246685. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 19.Wang X, Wang J, Zhang R, Huang Y, Feng S, Ma X, Zhang Y, Sikdar A, Roy R. Allelopathic effects of aqueous leaf extracts from four shrub species on seed germination and initial growth of Amygdalus pedunculata pall. Forests. 2018;9(11):711. 10.3390/f9110711. [Google Scholar]
  • 20.Hedge RS, Miller DA. Allelopathy and autotoxicity in alfalfa: characterization and effects of preceding crops and residue incorporation. Crop Sci. 1990;30(6):1255–9. 10.2135/cropsci1990.0011183X003000060020x. [Google Scholar]
  • 21.McNaughton SJ. Autotoxic feedback in relatin to germination and seedling growth in Typha latifolia. Ecology. 1968;49(2):367–9. 10.2307/1934475. [Google Scholar]
  • 22.Wang C, Liu Z, Wang Z, Pang W, Zhang L, Wen Z, Zhao Y, Sun J, Wang Z, Yang C. Effects of autotoxicity and allelopathy on seed germination and seedling growth in Medicago truncatula. Front Plant Sci. 2022;13:908426. 10.3389/fpls.2022.908426. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 23.Šoln K, Klemenčič M, Koce JD. Plant cell responses to allelopathy: from oxidative stress to programmed cell death. Protoplasma. 2022;259(5):1111–24. 10.1007/s00709-021-01729-8. [DOI] [PubMed] [Google Scholar]
  • 24.Abenavoli MR, Cacco G, Sorgona A, Marabottini R, Paolacci AR, Ciaffi M, Badiani M. The inhibitory effects of coumarin on the germination of durum wheat (Triticum turgidum ssp. durum, cv. Simeto) seeds. J Chem Ecol. 2006;32:489–506. 10.1007/s10886-005-9011-x. [DOI] [PubMed] [Google Scholar]
  • 25.Wang G, Qian J, Cheng G, Lan Y. Soil organic carbon pool of grassland soils on the Qinghai-Tibetan plateau and its global implication. Sci Total Environ. 2002;291(1–3):207–17. 10.1016/s0048-9697(01)01100-7. [DOI] [PubMed] [Google Scholar]
  • 26.Zhou X, Xiao Y, Ma D, Xie Y, Wang Y, Zhang H, Wang Y. The competitive strategies of poisonous weeds Elsholtzia densa benth. On the Qinghai Tibet plateau: allelopathy and improving soil environment. Front Plant Sci. 2023;14:1124139. 10.3389/fpls.2023.1124139. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 27.Han C, Pan K, Wu N, Wang J, Li W. Allelopathic effect of ginger on seed germination and seedling growth of soybean and Chive. Sci Hort. 2008;116(3):330–6. 10.1016/j.scienta.2008.01.005. [Google Scholar]
  • 28.Nishida N, Tamotsu S, Nagata N, Saito C, Sakai A. Allelopathic effects of volatile monoterpenoids produced by Salvia leucophylla: inhibition of cell proliferation and DNA synthesis in the root apical meristem of Brassica campestris seedlings. J Chem Ecol. 2005;31:1187–203. 10.1007/s10886-005-4256-y. [DOI] [PubMed] [Google Scholar]
  • 29.Đorđević T, Đurović-Pejčev R, Stevanović M, Sarić-Krsmanović M, Radivojević L, Šantrić L, Gajić-Umiljendić J. Phytotoxicity and allelopathic potential of Juglans regia L. leaf extract. Front Plant Sci. 2022;13:986740. 10.3389/fpls.2022.986740. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 30.Zeng R, Luo S, Shi Y, Shi M, Tu C. Physiological and biochemical mechanism of allelopathy of secalonic acid F on higher plants. Agron J. 2001;93(1):72–9. 10.2134/agronj2001.93172x. [Google Scholar]
  • 31.Bais HP, Vepachedu R, Gilroy S, Callaway RM, Vivanco JM. Allelopathy and exotic plant invasion: from molecules and genes to species interactions. Science. 2003;301(5638):1377–80. 10.1126/science.1083245. [DOI] [PubMed] [Google Scholar]
  • 32.Yildirim AN, Çelik C, Yildirim F, Şan B, Polat M, Binici S, Pepe AV. Effects of different plant growth-promoting sources on antioxidant enzyme activities and phenolic compounds of Apple. Appl Fruit Sci. 2025;67(3):1–9. 10.1007/s10341-025-01359-x. [Google Scholar]
  • 33.Sodaeizadeh H, Rafieiolhossaini M, Havlík J, Van Damme P. Allelopathic activity of different plant parts of Peganum harmala L. and identification of their growth inhibitors substances. Plant Growth Regul. 2009;59:227–36. 10.1007/s10725-009-9408-6. [Google Scholar]
  • 34.Gil CS, Duan S, Kim JH, Eom SH. Allelopathic efficiency of plant extracts to control cyanobacteria in hydroponic culture. Agronomy. 2021;11(11):2350. 10.3390/agronomy11112350. [Google Scholar]
  • 35.Wang Y, Hanson J, Mariam YW. Effect of sulfuric acid pretreatment on breaking hard seed dormancy in diverse accessions of five wild Vigna species. Seed Sci Technol. 2007;35(3):550–9. 10.15258/sst.2007.35.3.03. [Google Scholar]
  • 36.Wang C, Wu B, Jiang K. Allelopathic effects of Canada goldenrod leaf extracts on the seed germination and seedling growth of lettuce reinforced under salt stress. Ecotoxicology. 2019;28:103–16. 10.1007/s10646-018-2004-7. [DOI] [PubMed] [Google Scholar]
  • 37.Patanè C, Cavallaro V, Cosentino SL. Germination and radicle growth in unprimed and primed seeds of sweet sorghum as affected by reduced water potential in NaCl at different temperatures. Ind Crops Prod. 2009;30(1):1–8. 10.1016/j.indcrop.2008.12.005. [Google Scholar]
  • 38.Hou Q, Chen B, Peng S, Chen L. Effects of extreme temperature on seedling establishment of nonnative invasive plants. Biol Invasions. 2014;16:2049–61. 10.1007/s10530-014-0647-8. [Google Scholar]
  • 39.Li X, Wang J, Huang D, Wang L, Wang K. Allelopathic potential of Artemisia frigida and successional changes of plant communities in the Northern China steppe. Plant Soil. 2011;341:383–98. 10.1007/s11104-010-0652-3. [Google Scholar]
  • 40.Beauchamp C, Fridovich I. Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Anal Biochem. 1971;44(1):276–87. 10.1016/0003-2697(71)90370-8. [DOI] [PubMed] [Google Scholar]
  • 41.Zhou W, Zhao D, Lin X. Effects of waterlogging on nitrogen accumulation and alleviation of waterlogging damage by application of nitrogen fertilizer and Mixtalol in winter rape (Brassica napus L). J Plant Growth Regul. 1997;16:47–53. 10.1007/PL00006974. [Google Scholar]
  • 42.Quintanilla-Guerrero F, Duarte-Vázquez MA, García-Almendarez BE, Tinoco R, Vazquez-Duhalt R, Regalado C. Polyethylene glycol improves phenol removal by immobilized turnip peroxidase. Bioresour Technol. 2008;99(18):8605–11. 10.1016/j.biortech.2008.04.031. [DOI] [PubMed] [Google Scholar]
  • 43.Muñoz-Muñoz JL, García-Molina F, García-Ruiz PA, Arribas E, Tudela J, García-Cánovas F, Rodríguez-López JN. Enzymatic and chemical oxidation of trihydroxylated phenols. Food Chem. 2009;113(2):435–44. 10.1016/j.foodchem.2008.07.076. [Google Scholar]
  • 44.Aebi H. Catalase in vitro. In: Packer L, editor. Methods in enzymology. Orlando: Academic; 1984. pp. 121–6. 10.1016/S0076-6879(84)05016-3. [DOI] [PubMed] [Google Scholar]
  • 45.Brady CJ, Heng F. Rate of protein synthesis in senescing, detached wheat leaves. Funct Plant Biol. 1975;2(2):163–76. 10.1071/pp9750163. [Google Scholar]
  • 46.Arora R, Wisniewski ME, Scorza R. Cold acclimation in genetically related (sibling) deciduous and evergreen Peach (Prunus persica [L.] Batsch): I. seasonal changes in cold hardiness and polypeptides of bark and xylem tissues. Plant Physiol. 1992;105:95–101. 10.1104/pp.99.4.1562. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 47.Williamson GB, Richardson D. Bioassays for allelopathy: measuring treatment responses with independent controls. J Chem Ecol. 1988;14:181–7. 10.1007/BF01022540. [DOI] [PubMed] [Google Scholar]
  • 48.Turk MA, Tawaha AM. Allelopathic effect of black mustard (Brassica nigra L.) on germination and growth of wild oat (Avena fatua L). Crop Prot. 2003;22(4):673–7. 10.1016/s0261-2194(02)00241-7. [Google Scholar]
  • 49.Zhang T, Guo W, Tian X, Lv Y, Feng K, Zhang C. Allelopathic effects of Borreria latifolia on weed germination and identification of allelochemicals. J Sci Food Agric. 2025;105(1):626–34. 10.1002/jsfa.13859. [DOI] [PubMed] [Google Scholar]
  • 50.Haugland E, Brandsaeter LO. Experiments on bioassay sensitivity in the study of allelopathy. J Chem Ecol. 1996;22:1845–59. 10.1007/bf02028508. [DOI] [PubMed] [Google Scholar]
  • 51.Chon SU, Nelson CJ. Allelopathy in compositae plants. A review. Agron Sustain Dev. 2010;30(2):349–58. 10.1051/agro/2009027. [Google Scholar]
  • 52.Cruz Ortega R, Anaya AL, Ramos L. Effects of allelopathic compounds of corn pollen on respiration and cell division of watermelon. J Chem Ecol. 1988;14:71–86. 10.1007/BF01022532. [DOI] [PubMed] [Google Scholar]
  • 53.Bakkali F, Averbeck S, Averbeck D, Idaomar M. Biological effects of essential oils–a review. Food Chem Toxicol. 2008;46(2):446–75. 10.1016/j.fct.2007.09.106. [DOI] [PubMed] [Google Scholar]
  • 54.Arora S, Husain T, Prasad SM. Allelochemicals as biocontrol agents: promising aspects, challenges and opportunities. South Afr J Bot. 2024;166:503–11. 10.1016/j.sajb.2024.01.029. [Google Scholar]
  • 55.Mominul Islam AKM, Kato-Noguchi H. Plant growth inhibitory activity of medicinal plant Hyptis suaveolens: could allelopathy be a cause. Emirates J Food Agric. 2013;25(9):692–701. 10.9755/EJFA.V25I9.16073. [Google Scholar]
  • 56.Meychik N, Nikolaeva Y, Kushunina M. The significance of ion-exchange properties of plant root cell walls for nutrient and water uptake by plants. Plant Physiol Biochem. 2021;166:140–7. 10.1016/j.plaphy.2021.05.048. [DOI] [PubMed] [Google Scholar]
  • 57.Hu Y, Wang H. Production and scavenging of reactive oxygen species in plant cells under cadmium stress. Agron J. 2025;117(4):e70116. 10.1002/agj2.70116. [Google Scholar]
  • 58.Cheng F, Cheng Z. Research progress on the use of plant allelopathy in agriculture and the physiological and ecological mechanisms of allelopathy. Front Plant Sci. 2015;6:1020. 10.3906/bot-1302-29. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 59.Callaway RM, Ridenour WM, Laboski T, Weir T, Vivanco JM. Natural selection for resistance to the allelopathic effects of invasive plants. J Ecol. 2005;93(3):576–83. 10.1111/j.1365-2745.2005.00994.x. [Google Scholar]
  • 60.Wang L, Oduor AM, Liu Y. A native herbaceous community exerts a strong allelopathic effect on the Woody range-expander Betula fruticosa. J Plant Ecol. 2024;17(4):rtae055. 10.1093/jpe/rtae055. [Google Scholar]
  • 61.Wang C, Liu X, Li J, Yue L, Yang H, Zou H, Wang Z, Xing B. Copper nanoclusters promote tomato (Solanum lycopersicum L.) yield and quality through improving photosynthesis and roots growth. Environ Pollut. 2021;289:117912. 10.1016/j.envpol.2021.117912. [DOI] [PubMed] [Google Scholar]
  • 62.Wu H, Shabala L, Shabala S, Giraldo JP. Hydroxyl radical scavenging by cerium oxide nanoparticles improves Arabidopsis salinity tolerance by enhancing leaf mesophyll potassium retention. Environ Science: Nano. 2018;5(7):1567–83. 10.1039/C8EN00323H. [Google Scholar]
  • 63.Li D, Si J, Li J, Wang P, Yuan L. Physiological responses and differences of Populus euphratica to salt stress and drought stress. J Desert Res. 2023;43(2):205–15. 10.7522/j.issn.1000-694X.2023.00006. [Google Scholar]
  • 64.Rajput VD, Harish, Singh RK, Verma KK, Sharma L, Quiroz-Figueroa FR, Meena M, Gour VS, Minkina T, Sushkova S, Mandzhieva S. Recent developments in enzymatic antioxidant defence mechanism in plants with special reference to abiotic stress. Biology. 2021;10(4):267. 10.3390/biology10040267. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 65.Ding J, Sun Y, Xiao C, Shi K, Zhou Y, Yu J. Physiological basis of different allelopathic reactions of cucumber and Figleaf gourd plants to cinnamic acid. J Exp Bot. 2007;58(13):3765–73. 10.1093/jxb/erm227. [DOI] [PubMed] [Google Scholar]
  • 66.Hua Q, Liu Y, Yan Z, Zeng G, Liu S, Wang W, Tan X, Deng J, Tang X, Wang Q. Allelopathic effect of the rice straw aqueous extract on the growth of Microcystis aeruginosa. Ecotoxicol Environ Saf. 2018;148:953–9. 10.1016/j.ecoenv.2017.11.049. [Google Scholar]
  • 67.Gray WM. Hormonal regulation of plant growth and development. PLoS Biol. 2004;2(9):e311. 10.1371/journal.pbio.0020311. [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 68.Miransari M, Smith DL. Plant hormones and seed germination. Environ Exp Bot. 2014;99:110–21. 10.1007/s11258-012-0078-5. [Google Scholar]
  • 69.Oracz K, Voegele A, Tarkowská D, Jacquemoud D, Turečková V, Urbanová T, Strnad M, Sliwinska E, Leubner-Metzger G. Myrigalone a inhibits Lepidium sativum seed germination by interference with Gibberellin metabolism and apoplastic superoxide production required for embryo extension growth and endosperm rupture. Plant Cell Physiol. 2012;53(1):81–95. 10.1093/pcp/pcr124. [DOI] [PubMed] [Google Scholar]
  • 70.García M, García G, Parola R, Maddela NR, Pérez-Almeida I, Garcés-Fiallos FR. Root-shoot ratio and SOD activity are associated with the sensitivity of common bean seedlings to NaCl salinization. Rhizosphere. 2024;29:100848. 10.1016/j.rhisph.2024.100848. [Google Scholar]

Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

The dataset involved in this study are available with the following link: https://data.4tu.nl/private_datasets/3OjYjT4ODjjtxdnkXw2aGJr8fqUEBrkVsfiiThs93Pk.

Articles from BMC Plant Biology are provided here courtesy of BMC

RESOURCES