In nature, plants are often exposed to a variety of environments. The study of plant phenotypic plasticity cannot ignore a variety of environmental factors. At present, early exposure to flood or drought conditions will change the response of plants to later conditions, but little research has been done on whether early drought experience affects the interaction between plants in the later period. This paper takes Celtis sinensis and Bidens pilosa L. as the research object, through the second stage of individual growth ( drought, wet ), intraspecific interaction ( dry-dry, drought-wet, wet-wet ), interspecific interaction (dry-dry, drought-wet, wet-dry, wet-wet) biomass and morphological characteristics. The results showed that under the intraspecific interaction, the total biomass and aboveground biomass of Celtis sinensis owed a promoting effect under the treatment of no plasticity in the early flower pot ( no induced DP ) and no plasticity in the early stage and plasticity in the neighbor ( single induced SP- ), while the total biomass and aboveground biomass of Celtis sinensis showed a competitive effect under the treatment of plasticity in the early flower pot ( double induced NP ) and plasticity in the early stage and no plasticity in the neighbor ( single induced SP + ). It shows that early water-induced plasticity affects the strength of plant interaction in the later stage to varying degrees. It experienced early wetting ( no induced DP ). In the later intraspecific interaction, Celtis sinensis showed limited aboveground growth and good underground growth, which was contrary to the result that the aboveground part grew well and the underground part grew limited under single induced SP-. This indicates that the current plastic response of plants is not only dependent on early environmental experience, but also may be related to the cost of plasticity. Therefore, the early plastic response to drought environment may sacrifice the subsequent growth potential of Celtis sinensis and limit the plasticity of Celtis sinensis in the later stage. For Bidens pilosa L. under the intraspecific interaction, the aboveground biomass was increased and the underground biomass was decreased under the treatment of early plasticity and no plasticity of neighbors ( single induced SP + ), while the aboveground growth was limited and the underground growth was good under the treatment of early plasticity and plasticity of neighbors ( single induced SP- ). The influence of different early experiences on the interaction between early and late plants Different Bidens pilosa L. can cope with the influence of late intraspecific interaction through the mutual transformation of aboveground and underground parts.
Research Article
fects of early drought-induced phenotypic plasticity on late plant seedling interactions
https://doi.org/10.21203/rs.3.rs-5244739/v1
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phenotypic plasticity
early water experience
later intraspecific interaction
late interspecific interaction
biomass characteristics
Phenotypic plasticity is considered to be one of the main mechanisms for organisms to adapt to the environment[1 2]. Previously, most studies on phenotypic plasticity have considered the plasticity response to a single factor[3], usually non-biological factors (such as light, water, or nutrition). However, in the wild, plants tend to be exposed to multiple environments[4 5].Through field experiments in a more realistic environmental context and / or complex multi-factor experiments[6], multiple environmental factors cannot be ignored in the study of plant phenotypic plasticity. At present, there are related studies on the interaction between biological factors and non-biological factors[7]. For example, studies have shown that when different species interact, differences in their growth and adaptation strategies make them more favorable to cope with the lack of plant adaptation resources[8]. Plants have the ability to better respond to drought or wet conditions by adjusting local biomass when interacting with neighboring plants[6]. These studies take into account that plants respond to the interaction of multiple factors by regulating phenotypic plasticity. Some phenotypic results are significant, but some phenotypic results are not significant. The results of differentiation may be related to the direction of individual response and the direction of environmental impact in addition to biotic and abiotic factors[9]. However, the environmental background of plasticity and the response of organisms have certain complexity. It is difficult to separate the impact of plasticity from the treatment of plasticity from the confusion effect. It is difficult to effectively distinguish individual response from environmental impact (biological factors, non-biological factors). Distinguishing the two components of environmental impact and organism response is a prerequisite for effectively measuring the ability of organisms to respond to the environment.
Phenotypic plasticity is considered to be the main means for plants to cope with environmental changes. In the natural environment, in addition to being exposed to multi-factor environments, plants will also experience the possibility of being migrated to other environments. In the face of changes in this environment, plants can flexibly adapt to the environment by adjusting phenotypic plasticity. The ability of plants to produce plastic responses depends not only on the environment at the time, but also on early environmental experiences, such as priming or training, stress memory[10]. Ecological stress memory refers to that when the ecological events experienced by plants occur again, ecological memory will extract the previously accumulated experience, accelerate the system response speed, and enhance the system resistance, so as to complete the extraction and recognition process of ecological memory, so that it will have stronger stress tolerance in the future when facing stress environment[11 12]. Studies have shown that herbivores can induce defense responses in plant leaves when they eat plant leaves, thus affecting the performance of other herbivores. Changes in plant phenotypes due to prior damage have reduced interactions with subsequent antagonistic organisms[13], and studies have shown that. Transplanting plant communities to warmer climates may lead to increased competition with new competitors, that is, after plants migrate to new places, encounter new neighbors, and plant interactions may be changed. There have been related studies on early biologically induced plasticity, but there are few studies on the plasticity induced by abiotic factors. Therefore, the study of phenotypic plasticity should not only pay attention to the environmental background at the time of inducing plasticity, but also contact the early environmental events and habitat characteristics experienced by species[28]. The early environmental experience not only affects the phenotypic plasticity of plants, but also affects the future growth potential of plants. For example, the plastic response to shading in the early stage reduces the response of plants to shading in the later stage[14]; early drought or waterlogging has a significant effect on the response of plants to water in the later stage[25]. Plasticity research mainly focuses on the plastic response to the changes of current environmental factors, that is, the plasticity of plasticity, whether it is for abiotic factors or plant interactions. At present, the influence of early plant experience on plant interaction has not been explored.
In this paper, the effects of plasticity induced by early water experience on the strategy of plant interaction in the later stage were studied by using Celtis sinensis and Bidens pilosa L., which are common species in the pioneer weed community in karst habitats. Specifically answer the following questions: (1) Does early water induce plasticity? (2) Does early water treatment affect a) the strength of plant interaction at late stage ; b) Plant interaction-induced plasticity? (3) Whether there are differences in the effects on intraspecific and interspecific interactions.
3.1.1 Experimental materials
Celtis sinensis with good adaptability in karst limestone habitat was selected, which belongs to Ulmaceae. Bidens pilosa L., a common species of pioneer weed community in karst habitat, belongs to Asteraceae (Compositae). Pittosporum komarovii was collected from the same mother plant in the natural community of Shili River beach in Huaxi District, Guiyang City, Guizhou Province, and Bidens pilosa L. was collected from the same mother plant in the natural community near the experimental greenhouse in the western campus of Guizhou University.
3.1.2 Experimental design and processing
The experiment was carried out in the nursery of Forestry College, West Campus of Guizhou University, Huaxi District, Guiyang City, Guizhou Province. The site (106 ° 42'E, 26 ° 34'N) has an altitude of about 1020m, an average annual temperature of about 15.3°C, an average annual relative humidity of about 77%, an average annual total precipitation of about 1 129.5mm, an average annual rainy time of about 235.1d, an average annual sunshine hours of about 1148.3h, and an average annual snowfall time of 11.3d.
The experiment was conducted from July to October 2022 and lasted for 90 days. The seeds were placed in a tray and placed in an artificial climate incubator with temperature control and (25 ± 3) °C, humidity control at (60 % ± 2%), light and dark for 12 h each for seedling raising. Three weeks after germination, seedlings with the same growth were selected for transplantation into flowerpots ( inner diameter 22cm, depth 20cm ), and water treatment was started after one week of slow seedling. The test matrix formula was nutrient soil and quartz sand 1 ~ 1 mixed evenly.
The split-plot experimental design was used. The experiment was divided into two stages. In the first stage, environmental factors were treated to induce plant plasticity. The environmental factors were dry and wet, respectively. Species A and B were planted separately in pots, totaling 2 species × 2 water × 100 plants = 400. It was subjected to water treatment for 40 days. At the end of the first stage, destructive sampling was carried out, that is, 10 strains of A and B species under each water treatment were randomly selected, totaling 40 strains.
After the first stage of water treatment, the interaction strategy (single growth, intraspecific interaction, interspecific interaction) was performed. In order to make the plant roots fully contact to produce interaction, the plant spacing was less than 2cm. Each treatment was repeated 5 times, and the blocks under each treatment were repeated 3 times. For example, there are two species A and B in the first stage, drought is represented by −, and wetness is represented by +. The second stage of intraspecific interaction includes three early water treatments, namely drought-drought (A + and A +), drought-wet (A + and A-) and wet-wet (A- and A-) ; the second stage of interspecific interaction includes four early water treatments, namely drought-drought (A + and B +), drought-wet (A + and B-), wet-drought (A-and B +), wet-wet (A- and B-) (Figure 3.1).
In the second stage, under each interaction strategy treatment, in order to detect the effect of plants with early plastic response on plant interaction, the following is used as a marker. The first stage was subjected to drought treatment (with plasticity induction ) and wet treatment (without plasticity induction). In the second stage, there were three combinations under intraspecific interaction, including pots with plasticity treatment (double induction: NP), no plasticity treatment (no induction: DP), and only one plant in the same pot had plasticity : ( single induction: SP), where + indicated plasticity, that is, it had plasticity and its neighbors had no plasticity (single induction : SP +), -indicated plasticity, that was, it had no plasticity and its neighbors had plasticity treatment (single induction: SP-). (2 individual × 2 species + 3 intraspecies × 2 species × 2 plants + 4 interspecies × 2 plants) × 3 blocks × 5 replicates = 360 plants.
The first stage of treatment : environmental factors induce plant plasticity. Wet (+) and dry (-) water conditions were treated with species A and species B, respectively. The soil saturated water content was carried out according to the pre-experiment. Wet water was weighed every day, and water was weighed three days after drought. The other treatments are consistent. After 40 days of random destructive sampling, the second stage of interaction strategy treatment was carried out after the end of the first stage, and the seedlings were slowed down for 1 week. Keep the light, water and nutrition consistent, unified processing 40d.
3.1.3 Data collection and analysis
Before the first stage of treatment, the maximum leaf length of all plants was measured as the initial size. Destructive sampling was performed after the first stage lasted for 40 days, and the number of samples was 40. Before the beginning of the second stage, the maximum leaf length of the plant was measured again as the initial size of the second stage. The second stage lasted for 40 days and a series of morphological characteristics of all plants were harvested. For the 360 samples in the second stage, the morphological characteristics such as plant height, ground diameter, leaf area and leaf number were measured. Then the whole plant was sampled according to roots, stems and leaves, dried to constant weight in an oven at 85°C, weighed, and calculated total biomass, aboveground biomass, underground biomass and root-shoot ratio.
Plasticity analysis was performed on all the characteristics of the plants sampled after the end of the first stage. In order to explore the effect of plasticity on plant interaction in the second stage, (RDPIS, PI) was used to calculate the plasticity of a certain characteristic of a species. The formula is as follows :
X represents the average value of the adjusted characteristics under drought treatment, and Y represents the average value of the adjusted characteristics under wet treatment. The plasticity of species characteristics under drought treatment was significant at P<0.05 level.
For total biomass, aboveground and belowground biomass, the intraspecific and interspecific interaction intensity of each species was calculated using the relative interaction index (RII) interaction results[14]. The formula is as follows :
Among them: Bo represents the biomass of the target plant when growing alone, and Bw represents the biomass of the target plant when there are adjacent plants ( both from 0 to infinity ). The RII values range from-1 to 1, and the calculation results are positive to indicate promotion, and negative to indicate competition.
The relative growth (RG) of each trait of a plant that has undergone two rounds of treatment is calculated using the following formula:
Among them: X is the characteristic average value after early treatment, and Y is the characteristic average value of plants undergoing the same early treatment after the end of post-treatment. For example, X is the average total biomass of species after early drought, and Y is the average total biomass of species after early drought under late intraspecific interaction.
The experimental data were completed by IBM SPSS 2026 software and plotted by Origin 2018 software. All data were logarithmically converted before analysis to meet homogeneity of variance and normal distribution. After the end of the first stage, the effects of species and water treatment on each feature were analyzed by two-way ANCOVA analysis with two-way covariance analysis, and the effects of water treatment on all features were analyzed by one-way ANCOVA analysis with one-way covariance analysis.
After the end of the second stage, the effects of species, early treatment, and late species strategies on all features were analyzed by three-way ANCOVA with three-way covariance analysis, and one-way ANCOVA with one-way covariance analysis to analyze the effects of interaction strategies on all features. The total biomass was covariated with the initial maximum leaf length (initial size), and the other biomass and morphological characteristics were covariated with the total biomass. Multiple comparisons were performed using the least significance difference (LSD) method in the general linear model ( P<0.05 ).
3.2.1 Biomass and morphological characteristics responding to water conditions in the first stage of destructive sampling
After removing the individual size effect, species, water and their interaction had a significant effect on total biomass, and species had a significant effect on plant height (Table 3.1). Compared with wet (control) conditions, drought increased the biomass characteristics and root-shoot ratio of Celtis sinensis, that is, positive response (PI1.44 ~ 11.65) and increased plant height, ground diameter and leaf number. Drought only increased the root-shoot ratio and plant height of Bidens pilosa L., and reduced the total biomass, aboveground and underground biomass and other morphological characteristics (PI-0.05 ~ -0.24), that is the negative response (Figure 3.1).
Table 3.1 Two-way covariance analysis of the effects of species (SP), water conditions (W) and their interactions on all biomass and morphological characteristics in the first stage.
|
Source |
Df |
Lg10(TM) |
Lg10(AM) |
Lg10(UM) |
Lg10(R/S) |
Lg10(PH) |
Lg10(GD) |
Lg10(SLA) |
Lg10(LN) |
|
|
|
|
| ||||||
|
CO |
1 |
0.514 |
353.553*** |
71.692*** |
2.127* |
4.405 |
1.002* |
0.478 |
11.999** |
|
SP |
1 |
187.774*** |
1.614 |
1.614 |
1.123 |
2.637* |
0．074 |
0.123 |
6.333 |
|
W |
1 |
0.906*** |
1.943 |
1.943 |
3.087 |
2.005 |
1.003 |
9.537 |
5.194 |
|
SP×W |
1 |
32.119*** |
1.805 |
1.805 |
0.48 |
0.12 |
0.227 |
0.538 |
1.678 |
Note: The initial size (the length of the maximum leaf ) was the covariate for total biomass , and total biomass was the covariate for all the other traits. *P<0.05, **P<0.01, ***P<0.001.
3.2.2 Effects of early plasticity on plant late performance and response
After removing the individual size effect, species, early and late treatments and their interactions all had significant effects on most traits (Table 3.2). Under the intraspecific interaction, the early self-plasticity and neighbor plasticity ( single induction: SP- ) treatment increased the total biomass, aboveground biomass, plant height and ground diameter (ANCOVA, LSD, P<0.05), and decreased the underground biomass, root-shoot ratio and leaf number (ANCOVA, LSD, P<0.05). However, in the early pots, both of them had no plasticity (no induction: DP) treatment significantly reduced the total biomass, aboveground biomass, plant height and ground diameter of Celtis sinensis (ANCOVA, LSD, P<0.05) ; the underground biomass, root shoot ratio and leaf number were increased (ANCOVA, LSD, P<0.05). For Bidens pilosa L., only the early self-plasticity and neighbor non-plasticity (single induction: SP +) treatment reduced the total biomass, underground biomass and plant height (ANCOVA, LSD, P<0.05), and increased its aboveground biomass and ground diameter. On the contrary, the early self-plasticity and neighbor plasticity ( single induction: SP-) treatment reduced the aboveground biomass of Bidens pilosa L. and increased its underground biomass, plant height and ground diameter ( ANCOVA, LSD, P< 0.05). The other three early treatments had no significant difference in the aboveground and underground biomass of Celtis sinensis.
Under interspecific interaction, no plasticity ( no induction: DP ) treatment in early flowerpots significantly reduced the root-shoot ratio of Celtis sinensis, and no plasticity in early stage and plasticity in neighbors ( single induction: SP- ) treatment significantly increased the root-shoot ratio ( ANCOVA, LSD, P<0.05 ). Early pot plasticity ( double induction: NP ) treatment and no plasticity ( no induction: DP ) treatment reduced the ground diameter and leaf number (ANCOVA, LSD, P<0.05), respectively. Early pot plasticity ( double induction: NP ) treatment reduced the ground diameter and increased the number of leaves ( ANCOVA, LSD, P<0.05 ), which was contrary to the results of early self-plasticity and neighbor non-plasticity ( single induction: SP +) treatment ( Fig.3.2 ; 3.3 ).
Table 3.2 Ternary covariance analysis of the effects of species (SP), early treatment (EW), late treatment (LI) and their interactions on morphological and biomass characteristics in the second stage.
|
Source |
Df |
Lg10(TM) |
Lg10(AM) |
Lg10(UM) |
Lg10(R/S) |
Lg10(PH) |
Lg10(GD) |
Lg10(SLA) |
Lg10(LN) |
|
|
|
|
|
|
|
| |||
|
CO |
1 |
11.847*** |
3903.911*** |
512.511*** |
.012*** |
19.132*** |
3.997*** |
5.894** |
22.146*** |
|
SP |
1 |
135.204*** |
9.637** |
9.637*** |
2.223*** |
511.322*** |
35.451*** |
3.734*** |
46.888*** |
|
EW |
1 |
6.181*** |
9.718** |
9.718** |
.581** |
2.603 |
.284*** |
.013 |
54.272*** |
|
LI |
2 |
6.402** |
6.610** |
6.610* |
.563* |
.500 |
2.432 |
.274 |
3.799* |
|
SP ×EW |
1 |
103.080*** |
.024 |
.024*** |
.854*** |
14.048** |
17.571*** |
1.183*** |
10.740** |
|
SP×LI |
2 |
20.562*** |
.611 |
.611* |
.179 |
1.303 |
.920 |
.546 |
11.110*** |
|
EW×LI |
2 |
5.727** |
3.407* |
3.407* |
1.000* |
.310 |
1.293 |
.290 |
2.539 |
|
SP×EW×LI |
2 |
11.060*** |
.378 |
.378 |
.866 |
.574 |
2.853 |
3.621*** |
1.778 |
Note: The total biomass takes the initial size (maximum leaf length) as the covariate, and other characteristics take the total biomass as the covariate. *P< 0.05, **P< 0.01, ***P< 0.001.
3.2.3 Effects of early plasticity on plant interaction strength
Under the intraspecific interaction, the total biomass and aboveground biomass of Celtis sinensis showed a promoting effect under the treatment of no plasticity ( no induction : DP ) in the early flowerpot, and its aboveground biomass showed a promoting effect under the treatment of no plasticity of itself and plasticity of its neighbors ( single induction : SP- ) in the early flowerpot. The total biomass and aboveground biomass showed a competitive effect under the early self-plasticity and neighbor non-plasticity ( single-induced SP + ) treatment and the early pot plasticity ( double-induced : NP ) treatment.
Under the interaction between species, the total biomass and aboveground biomass of Bidens pilosa showed a competitive effect under the treatment of no plasticity ( no induction: DP ) in the early flowerpot, but showed a promoting effect under the treatment of other early three ( double induction : NP, single induction: SP +, single induction: SP- ) ( Fig.3.4 ).
3.2.4 Relative growth of biomass and morphological characteristics of the two plants
Compared with the early wetting, the early self-plasticity and neighbor plasticity ( single induction : SP- ) and both non-plasticity ( no induction : DP ) treatments increased the total biomass and aboveground biomass of Celtis sinensis under the late intraspecific interaction, and increased the underground biomass and root-shoot ratio of Bidens pilosa L. under the late intraspecific interaction. Compared with early wetting, early self-plasticity and neighbor plasticity ( single induction: SP- ) treatment increased the total biomass, aboveground and underground biomass of Bidens pilosa under late interspecific interaction ( P < 0.05 ).
Compared with growing alone, intraspecific interactions increased the underground biomass and leaf number of Celtis sinensis under the treatment of no plasticity ( no induction: DP ) in early flowerpots. Interspecific interactions increased the total biomass, aboveground biomass and leaf number of Bidens pilosa L. under the treatment of plasticity ( double induction: NP ) in flowerpots ( P < 0.05 ) ( Fig.3.5 ; Fig.3.6 ).
3.3.1 Effects of early plasticity on plant interactions
Abiotics can change the results of interactions between plants. It is generally believed that competition between plants is more common under sufficient resources. When the environmental resources are relatively scarce, the competition is weakened or the promotion effect is enhanced, and the intensity of the promotion effect will increase with the severity of the environment[15]. This hypothesis is supported by follow-up studies. However, it is unknown whether the lack of early experience plasticity can change the later plant-plant interaction. The results of this study showed that under the intraspecific interaction, Celtis sinensis showed a promoting effect under the early self-plasticity and neighbor plasticity (single induction: SP-) treatment and early flowerpot plasticity (no induction: DP) treatment, while it showed a competitive effect under the other two early treatments. The plasticity induced by early experience led to different competitive abilities of Celtis sinensis and neighboring plants. Studies have shown that The results of the interaction depend on the stress intensity. In the results of this study, because the experiment is divided into two stages, the results of the interaction between plants in the later stage not only depend on the early environmental experience, but also may be related to the intensity of competition and species. When two species interact, a species' own plasticity is stronger, plasticity is conducive to promoting the occurrence of effects, on the contrary, it is more prone to competitive effects. In the early stage, Bidens pilosa L. showed a promoting effect under the treatment of pot plasticity (double induction: NP) and its own plasticity and no plasticity of neighbors (single induction: SP +). In the early stage, it showed competition under the treatment of no plasticity and plasticity of neighbors (single induction: SP-). The competition intensity of early environmental experience also affected the growth potential of plants in the later stage, and further affected the results of interaction between plants in the later stage.
Under the treatment of no plasticity in the early stage and plasticity in the neighbors (single induction: SP-) and no plasticity in the flowerpot (no induction: DP), the intraspecific interaction of Celtis sinensis was more likely to promote the effect than the interspecific interaction. In the early stage, it experienced a humid environment, and the demand for water between the same species in the later stage was similar, which alleviated the excessive water and promoted the growth of plants. The different environmental experiences of the adjacent plants in the early stage also made the aboveground and underground parts of Celtis sinensis different. The results of Bidens pilosa L. and Celtis sinensis are opposite. Bidens pilosa L. has less water demand than Celtis sinensis, and has experienced a humid environment in the early stage. In the later stage, the underground part of Bidens pilosa L. grows well to alleviate the competition of the aboveground part. Studies have shown that the intraspecific competition effect of the same neighbor is more significant because the niche overlap of the same neighbor is higher than that of the different neighbor, and the way of obtaining resources is more similar.
3.3.2 Effects of early plasticity on plant late performance and response
Previously there was a response to moisture [18]or plant-plant interactions [19], but rarely based on a certain stage of plant life history to explore related issues. The results of this study show that under intraspecific interactions, early self-plasticity and neighbor plasticity ( single induction : SP- ) treatment increased the total biomass and aboveground biomass of Celtis sinensis, and reduced underground biomass and root-shoot ratio. This means that the early experience of moist (single induction: SP-) treatment, the late intraspecific interaction showed that the growth of the aboveground part was good and the growth of the underground part was limited. According to the optimal allocation theory, plants should allocate resources to the organs that obtain the most limited resources[20]. For example, plants will allocate more biomass to the roots proportionally under water-deficient conditions to obtain more water[21]. In the later period, the interaction within the species promoted the aboveground part and inhibited the underground part, which may be due to the fact that the adjacent plants experienced the drought environment in the early stage, resulting in the underground part growing well. In the later period, due to the same demand for resources among the same species[22], the underground part of the adjacent plants ( neighbors with plasticity ) experiencing drought is better, which limits the growth of the underground part of their own plants. The final result is that the increase of aboveground biomass is at the expense of underground biomass. Studies have shown that the root ' tragedy of the commons ' research results show that the increase of intraspecific competitive root biomass is at the expense of the reproductive biomass of the plant[23 24], the presence of the same neighbor can induce additional stem elongation at the expense of reduced leaf growth[25], which explains why the growth of the aboveground part is good and the growth of the underground part is limited.
Studies have shown that early drought can improve the biomass growth of plants in the late flooding environment[26]. In this study, the early wetting ( no induction: DP ) was experienced, and the later intraspecific interaction showed that the aboveground growth was limited and the underground growth was good. The results of this study are contrary to the above results that the aboveground part grows well and the underground part is limited under single induction SP-, which once again proves that the results of the aboveground and underground parts are not only related to their own early experience of water, but also related to the early experience of neighboring plants. In both cases ( single induction : SP-and non-induction : DP ), the interaction with the same species occurred in the later stage. The only difference was that the neighboring plants experienced different water conditions in the early stage. The neighboring plants under single induction SP-were affected by drought conditions, while the neighboring plants under non-induction DP were affected by wet conditions, indicating that the intraspecific interaction showed different plasticity in the later stage, which was affected by the early water experience of neighboring plants. For Bidens pilosa L., under the intraspecific interaction, the aboveground biomass was increased and the underground biomass was decreased under the treatment of early plasticity and no plasticity of neighbors ( single induction: SP + ), while the aboveground growth was limited and the underground growth was good under the treatment of early plasticity and plasticity of neighbors ( single induction: SP- ). It is shown again that the opposite results under the second-stage plant intraspecific interaction strategy are related to the early water experience. When the other two species had the same early experience, the aboveground and underground parts showed the opposite situation, which is related to the species strategy.
Whether for itself or neighboring plants, the results of these differences may be related to the early water experience, and may also be related to the early effect on the later growth potential. The early plastic response to the humid environment may sacrifice the subsequent growth potential of Celtis sinensis, thus limiting the plasticity of Celtis sinensis in the later stage. Studies have shown that the response of plants to the current environment depends on the early plastic response, that is, the expression of traits in the early stage of plant ontogeny may affect the expression of traits in the later stage of ontogeny, and there will be historical limitations[27]. Studies have shown that early exposure to flood or drought conditions can change the plant 's response to later conditions and increase the tolerance to extreme conditions in the later period[26]. The results of this study showed that the aboveground part grew well under single induced SP-treatment, and the growth of the underground part was limited. The opposite results were observed under non-induced DP treatment. One explanation may be that the long-term growth of plants under high water conditions will cause waterlogging stress, during which the inhibition of root respiration and the accumulation of toxic substances will affect the production of plant biomass[29]. Another explanation is that the later stress environment may increase the cost[30], which is considered to be a ' life history limitation' of plasticity. The later positive or negative plasticity response is not only related to the cost of plasticity, but also may be related to the later species strategy. Therefore, when plants are adversely affected by the external environment, plants can flexibly adjust their own plasticity according to the environment and the habits of neighboring plants, and can better adapt to environmental changes through the regulation of phenotypic plasticity.
3.3.3 Effects of early plasticity on plant responses to plant-plant interactions
Compared with individual growth, intraspecific interaction was more likely to increase the aboveground biomass of Celtis sinensis under its own plasticity and neighbor non-plasticity treatment ( single induction: SP + ), and reduce its underground biomass and root-shoot ratio. In the same early stage, the interspecific interaction was more likely to reduce the aboveground biomass and increase the underground biomass of Celtis sinensis under the treatment of plasticity and no plasticity of neighbors ( single induction: SP + ). It shows that the results of interspecific interaction and intraspecific interaction are opposite when subjected to the same water experience in the early stage. The above-ground and below-ground interactions in intraspecific and interspecific interactions were still the opposite results under the treatment of non-plasticity in itself and plasticity in neighbors ( single induction: SP- ), which indicated that the results of the second stage were not only affected by the later neighboring plants, but also showed that the later stage under the same species strategy. When experiencing different water environments in the early stage, the performance of the above-ground and below-ground in the later stage was also the opposite, and the same species in the later stage. In the case of species interaction, a plant itself experiences drought environment in the early stage and adjacent plants experience wet environment interaction in the early stage. Due to early exposure to two opposite water environments of drought and wet, the early plastic response to wet conditions may sacrifice the growth potential of plants in the later stage, and the later pressure environment may increase the cost of plasticity[30].This ' life history limitation ' is manifested in late growth limitation. When experiencing the same water environment in the early stage, the aboveground and underground parts showed opposite results when interacting with the same or different species in the later stage, which was consistent with the results of plasticity studies in response to intraspecific and interspecific interactions.
For Bidens pilosa L. intraspecific interactions were more likely to increase the above-ground biomass and reduce the below-ground biomass under the plasticity treatment ( double induction : NP ) of the flowerpots compared with the single growth, which was the same as the results of interspecific interactions ; intraspecific interaction was more likely to increase aboveground biomass and reduce belowground biomass under the treatment of plasticity in itself and no plasticity in neighbors ( single induction: SP+ ). Interspecific interaction was more likely to increase aboveground biomass and reduce belowground biomass under the treatment of both plasticity in flowerpots ( no induction: DP ). Whether it is affected by different early water or later interaction with different adjacent plants, the final performance of Bidens pilosa L. is that the ground grows well and the underground growth is limited, which once again shows that the early water experience is different but the later performance is consistent. However, whether B.pilosa experienced different early water environments or interacted with the same or different neighboring plants in the later stage, the aboveground and underground showed opposite growth phenomena, which may be related to species specificity and further affect the plasticity response.
This study shows that early water-induced plasticity can change the results of plant interaction in the later stage, and the plasticity response of plant interaction in the later stage is different. The results of plant-plant interactions in the later stage not only depend on the early environmental experience, but also may be related to the intensity of competition and species. Under the treatment of no plasticity of itself and plasticity of its neighbors ( single induction : SP- ) and no plasticity of both ( no induction: DP ), the intraspecific interaction of Celtis sinensis is more likely to induce the promotion effect than the interspecific interaction, and Bidens pilosa L. is the opposite of this result. This may be related to the intensity of competition and species specificity caused by early environmental experiences. Under the intraspecific interaction, the treatment with no plasticity and plasticity of neighbors ( single induction: SP- ) promoted the aboveground growth of Celtis sinensis and inhibited the underground growth, which was contrary to the results of both treatments without plasticity ( no induction : DP ). This may be related to the early water-induced plasticity. It can be seen from the late morphological characteristics of Celtis sinensis that the current plastic response of plants not only depends on the early environmental experience, but also may be related to the cost of plasticity. Therefore, the early plastic response to arid environment may sacrifice the subsequent growth potential of Celtis sinensis and limit the plasticity of Celtis sinensis in the later stage. For Bidens pilosa L. under intraspecific interactions, early self-plasticity and neighbor plasticity ( single induction: SP + ) treatment promoted the growth of aboveground parts and limited the growth of underground parts, which was contrary to the results of early self-plasticity and neighbor plasticity ( single induction: SP- ) treatment. Bidens pilosa L. can cope with the influence of later intraspecific interactions through the mutual transformation of aboveground and underground parts. When the early experience changed, the aboveground and underground parts of Bidens pilosa L. showed completely opposite results, and the biomass characteristics of intraspecific and interspecific were not significantly different, indicating that it was not only related to plant strategies, but also the early environmental experience could affect the later growth potential.
Author Contribution
Hou Xiali completed data analysis and article writing. Teacher Wang Shu completed the article to guide writing and inspection Yang Qingzhu and Huang Yuanhu helped to complete the experimental operation.
Acknowledgement
Ethics approval and consent to participateAll methods were performed in accordance with the relevant guidelines and regulations of institutional, national, and inter national guidelines and legislatConsent for publicationNot applicable.Clinical trial number: not applicableAvailability of data and materialWilling to provide the original data and open, the original data has been uploaded to the attachment.FundingThis research was provided by the National Natural Science Foundation of China (NSFC, 32171511)Competing interestsWe declare that there is no competing interest among the authors.AcknowledgementsWe are grateful for the reviewers and editors who all provided useful feedback on this manuscript.
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No competing interests reported.