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Article

Is Clonal Integration a Buffer for the Stress of Resource Acquisition Depletion in Eichhornia crassipes (Pontederiaceae) Ramets?

by
Guilherme Ramos Demetrio
1,2,*,
Dalton Serafim
2 and
Flávia de Freitas Coelho
3
1
Plant Ecology Lab, Unidade Educacional Penedo, Campus Arapiraca, Federal University of Alagoas, Penedo 57200-000, Alagoas, Brazil
2
Pós-Graduação em Diversidade Biológica e Conservação nos Trópicos, Instituto de Ciências Biológicas e da Saúde, Campus A. C. Simões, Universidade Federal de Alagoas, Maceió 57072-970, Alagoas, Brazil
3
Departamento de Biologia, Universidade Federal de Lavras, Lavras 37203-202, Minas Gerais, Brazil
*
Author to whom correspondence should be addressed.
Stresses 2024, 4(4), 734-743; https://doi.org/10.3390/stresses4040047
Submission received: 10 October 2024 / Revised: 28 October 2024 / Accepted: 31 October 2024 / Published: 2 November 2024
(This article belongs to the Collection Feature Papers in Plant and Photoautotrophic Stresses)
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Abstract

Natural selection favors the allocation of finite resources to different functions maximizing fitness. In this sense, some functions may decrease whereas others increase when resources are limited in a process called a trade-off. However, a great variety of situations may obscure trade-off detection in clonal plants, such as the ability to generate offspring by clonal growth that represents opportunities for resource uptake. The aim of this work was to evaluate if clonal integration and resource availability mediate biomass allocation patterns in E. crassipes through a greenhouse experiment. We set ramets in clonal and isolated conditions, and with and without leaf blades, and compared the relationship of biomass proportion allocated to each vegetative organ. We found that biomass allocation to vegetative structures in E. crassipes is primarily shaped by resource pools and is enhanced by clonal integration as attached ramets invest more in growth and vegetative structures. In this sense, regarding trade-off patterns in biomass allocation among vegetative organs and under resource depletion, clonal integration may represent a way to stabilize biomass allocation patterns and may decrease trade-off importance. We discuss trade-offs and clonal integration as evolutionary strategies that allow plant persistence and improve plants fitness. These findings may support aquatic plant management and control efforts while highlighting the evolutionary significance of clonal integration for plant life strategies.

1. Introduction

Plants thrive in diverse ecosystems, which each present unique environmental conditions that often impose stressful conditions, such as nutrient scarcity or drought [1,2]. These situations challenge plants’ survival by limiting the availability of essential resources, such as mineral nutrients and solar radiation, required for metabolic processes. These resource constraints force plants to optimize their allocation strategies, prioritizing organs that maximize survival and growth under the prevailing conditions. Since plants depend on finite environmental resources to support their metabolic functions, they must efficiently allocate these resources to the most essential organs for growth and survival in each moment of an individual’s life history [3]. This allocation becomes particularly important under stress, where limited resources can accentuate trade-offs between different vegetative organs. Therefore, resource allocation plays a crucial role in shaping plant life histories and their responses to environmental change.
Optimal biomass partitioning theory [4] postulates that plants allocate more resources to organs that acquire the most limiting resource, while life-history theory [5] suggests that plants prioritize organs essential for survival or reproduction to maximize fitness over time. These variations in allocation dynamics among different organs are key to understanding phenotypic plasticity [6] and, consequently, the ecological strategies employed in response to environmental conditions [7]. Thus, a plant’s success depends on how it allocates biomass among roots, stems, leaves, and flowers in response to environmental conditions, especially under stress.
Under severe resource limitation, plants may prioritize certain functions or organs at the expense of others, leading to trade-offs in biomass allocation [8]. For example, plants under stress, such as drought, may present a greater allocation of biomass to roots as a way of increasing water acquisition [9,10]. Additionally, resource competition [11] and nutrient stress [12] also drive altered patterns of biomass allocation. Defoliation is another stress that can cause drastic variations in plant biomass allocation. The removal of leaves creates situations of resource acquisition limitation since it reduces the photosynthetic area of the plant [13,14]. In addition, defoliation can also generate trade-offs in the allocation of biomass between different plant organs. For example, after defoliation, plants can decrease the allocation of biomass to stems to deal with carbon scarcity, increasing the investment of this nutrient in the construction of new leaves [15,16]. This pattern reflects a trade-off between carbon sink organs and carbon-synthesizing organs [16]. In this sense, trade-offs are a common process of plant ecology, which are often observed when plants face reproductive challenges or resource competition.
Trade-offs in resource allocation have been well-documented in plant ecology, particularly concerning reproductive strategies [17,18,19,20,21]. However, in clonal plants, where integration allows resource sharing between connected ramets, the severity of these trade-offs might be reduced because clonal integration could lead to a more flexible response to stress by redistributing resources more evenly across ramets [22], thereby potentially smoothing trade-offs that occur in stressed environments. Thus, clonality may also bias biomass allocation patterns since clonal plants may function as highly integrated units [23,24]. This is especially true regarding aquatic plants, such as Eichhornia crassipes (Mart.) Sölms. (Pontederiaceae), in which the extensive ability to generate offspring by clonal growth [25,26] may represent an opportunity for new resource acquisition and/or storage organ generation [27].
Building on this theoretical framework, we aimed to: (i) test if clonal integration influences biomass allocation patterns in Eichhornia crassipes, and (ii) assess whether clonal integration mitigates trade-offs among vegetative organs under stressful conditions of resource limitation. We hypothesized that clonal integration would buffer the effects of resource limitation, alleviating trade-offs and facilitating a more even allocation of biomass between vegetative organs compared to plants isolated from parental ramets.

2. Results

Among all measured traits, ‘defoliation’ treatment was the only one that significantly affected biomass allocation patterns. Defoliated ramets that remained attached to their parental plants exhibited lower biomass compared to other treatments. The significance and direction of the interactions among treatments and biomass allocation patterns varied for each variable, with ‘defoliation treatment’ being the only one that notably altered the slopes of relationships.
The proportion of biomass allocated to stems did not change in response to biomass allocation to the shoot components of ramets. Additionally, the proportion of biomass allocated to leaf blades (F = 1.89, p = 0.1092) and the proportion allocated to petioles (F = 1.96, p = 0.09) also showed no significant effect on biomass allocation to stems.
The proportion of biomass allocated to the root system decreased as the biomass allocation to aerial parts increased (Figure 1A,B). This decrease was more pronounced when the response variables were related to the shoot components of the plant (Table 1). The proportion of biomass allocated to stems exhibited a negative effect, albeit less intense, and showed no interaction with the applied treatments. Regarding shoot components, the proportions of biomass allocated to leaf blades and petioles decreased as the allocation to stems or roots increased (Table 1). In both cases, the allocation to roots had a more pronounced effect on the allocation to shoot parts (Table 1).
In Table 1, the results reveal distinct biomass allocation patterns among the vegetative organs of Eichhornia crassipes under different treatment conditions. The proportion of biomass allocated to roots decreased significantly as the allocation to shoot components (stems, leaf blades, and petioles) increased. This pattern was particularly influenced by defoliation treatment, indicating a strong reallocation response under resource limitation. Furthermore, while defoliated, isolated ramets showed a notable decrease in root biomass proportion; connected ramets under clonal integration displayed a more balanced root-to-shoot allocation, which was likely due to resource sharing facilitated by clonal integration.
Figure 1A illustrates how biomass allocation between roots and leaf blades varied with clonal integration and defoliation. In attached ramets, root-to-leaf blade allocation remained relatively stable despite defoliation, suggesting that clonal integration buffers the effects of leaf area loss. By contrast, isolated, defoliated ramets showed a steeper decline in root biomass allocation relative to leaves, indicating a prioritization of above-ground structures under resource scarcity to enhance light capture and potential carbon gain. Figure 1B,C further support the role of clonal integration in maintaining stable biomass allocation patterns. In Figure 1B, root-to-petiole allocation patterns shift markedly with defoliation, with isolated ramets showing a pronounced reduction in root investment compared to clonal treatments. Figure 1C reveals a positive correlation between biomass allocated to leaf blades and petioles, especially in clonal treatments, highlighting that clonal integration can stabilize resource distribution across shoot parts despite defoliation. These patterns reinforce that clonal integration buffers trade-offs between root and shoot allocation under stress.
Regarding aerial parts—leaf blades and petioles—the interaction term between the proportion of biomass allocated to each response variable and the ‘defoliation’ treatment was significant. This interaction revealed a smoother slope in the decrease of biomass allocation to leaf blades and petioles compared to clonal treatment (Table 1; Figure 1A,B).
For most treatments, except for ‘defoliation’ treatment, leaf blades and petioles did not follow the pattern of trade-off found for the other measured traits. These organs’ biomass showed a positive and significant correlation, indicating an influence of the interaction between clonality and defoliation for these traits’ biomass allocation (Table 1) (Figure 1C).

3. Discussion

As expected, our results show that E. crassipes biomass allocation is mediated by trade-offs occurring between root and shoot parts, and these trade-offs are buffered by clonal integration when the ramet remains linked to its mother plant. Additionally, stress seems to be a potential driver of biomass accumulation since decreases in resource availability (represented in this study by resource depletion caused by defoliation) may cause an alteration in how E. crassipes ramets allocate biomass to its vegetative organs.
Differences in allocation to structures and, therefore, functions are probably more important to ecological questions than differences in physiological mechanisms occurring at cellular or lower levels [28] since biomass allocation is closely related to the survival and growth strategy of plants [29,30]. This may occur mainly because phenotypic plasticity allows a genome to modify growth and development in response to changes in the environment [31], which may lead individuals that are adjusted to their habitats to show trade-offs related to environmental conditions [32]. For example, defoliation and natural herbivory can significantly influence biomass allocation in aquatic macrophytes [33]. In response to this resource depletion, macrophytes may reallocate resources from above-ground to below-ground structures, or vice versa, with species-specific responses. This reallocation is closely tied to the extent of leaf area loss and the availability of environmental resources. Eichhornia crassipes, for instance, compensates for low defoliation levels by continually investing in leaf production, though this response diminishes under higher levels of simulated defoliation [34]. Our data support these ideas as defoliation changed the allocation patterns for almost every organ we measured in our experiment, except for stems.
The presence of trade-offs at the ramet level in aquatic macrophytes remains a subject of debate in the literature. For instance, Eichhornia crassipes showed no trade-offs in biomass allocation to vegetative organs in response to density, and thus, competition in an Amazonian river [35]. A similar pattern was observed for Pistia stratiotes in the Pantanal [36]. These studies suggest that the high morphological plasticity of aquatic macrophytes and the role of clonal integration may mitigate trade-offs [24]. On the other hand, other studies found trade-offs for root and shoot allocation [37,38]. In cases where trade-offs are present, they are commonly linked to important aspects of plant fitness, such as nutrient assimilation by roots, and leaf investment [39]. This holds true for our dataset since we found that a major part of relationships among E. crassipes vegetative organs is mediated by trade-offs, except for the relationship between biomass allocation to leaf blades and petioles, which was positive.
This is a clear pattern when resource acquisition strategies are considered. Clonal plants use these strategies in response to resource allocation among ramets and adjust to particular environmental changes [40]. For the genus Eichhornia, previous works have shown that for rooted species, like E. azurea, nutrient level is an important driver for plant biomass allocation, with plants located in lentic habitats showing a strong trade-off among vegetative characteristics. This is due to lower water renewal in this kind of habitat, which is a characteristic found in our study, conducted in pots under greenhouse conditions. Water flow promotes water renewal [41], which avoids nutrient depletion in a variety of habitats colonized by aquatic plants. This absence of nutrient renewal through water movement may be of extreme importance for floating plants because aquatic habitats tend to be highly homogeneous [25], and nutrients tend to descend on the water column, becoming unavailable for root uptake.
In the case of E. crassipes, the stem is just the connective part between root and shoot parts, with the function of anchoring leaves and roots. Not surprisingly, the proportion of plant total biomass allocated to this organ was not different between resource levels, nor different clonal integration conditions. Root traits, however, are very important to evaluate the degree of plant adjustment to new environmental conditions [42], and changes in root morphology are commonly observed in aquatic plants, with roots being bigger in habitats that show higher stress levels, such as high plant density [35], low nutrient levels [36], or increasing size in the gradients of aquatic to terrestrial habitats [37]. In our dataset, roots decreased in biomass when an increase in leaves biomass was observed mainly due to the resource limitation to which plants were submitted. In this situation of resource limitation, this trade-off indicates a more pronounced reallocation of biomass from roots to leaves, which suggests an optimization of light resource acquisition [43] that may increase carbon assimilation rates in situations of defoliation stress. It is also important to note that this pattern was most pronounced in situations where resources could not be transported from parent ramets to offspring.
In this sense, clonal integration, under severe resource limitation, may decrease the importance of trade-offs, allowing a more balanced root/shoot ratio during daughter ramets growth. Therefore, clonal integration is advantageous to the clonal growth of parents and offspring ramets as whole [44], which is corroborated by changes in the whole plant in a way that enhances the uptake of the most limiting resource when all ramets lie within a homogeneous patch of habitats [45]. For our data, severed ramets showed a different response to clonal integration when compared to those that remained intact but not when isolated. This demonstrates the importance of clonal integration on translocating resources and allowing ramets’ plastic response as occurred for Hydrocotyle bonariensis in which severed ramets showed more variable responses to nutrient levels [46].

4. Materials and Methods

4.1. Study Species

Eichhornia crassipes (Mart.) Sölms., which is commonly known as water hyacinth, is a free-floating aquatic macrophyte [47,48] that is native to the Amazon River Basin [48]. In recent decades, it has gained recognition as an aggressive invasive species [49], and now exhibits a global distribution largely driven by its ornamental value [50]. The plant is characterized by a short stem from which leaves emerge in a rosette arrangement along with several fine roots [48]. Additionally, E. crassipes reproduces vegetatively through rhizome sprouting, allowing it to rapidly cover vast water surfaces [51]. Clonality seems to be an important life-history trait for this species since previous studies showed a significant effect of clonal integration with sexual reproductive traits, and clonal ramets exhibited larger and more robust flowers than isolated ones [24].

4.2. Plant Sampling

All plant material was collected from a large monospecific mat located at Represa do Funil (21°8′36.45″ S, 45°2′11.12″ W, Lavras, MG, Brazil). In the field, we sampled and washed the plants to remove debris. We selected adult ramets, identified by the presence of newly produced sexual reproductive structures or remnants of old floral scapes, with no visible signs of foliar herbivory or disease. A total of 90 ramets were selected, placed in plastic bags with a small amount of water to prevent root desiccation, and transported to a greenhouse at the Agronomy Department of Federal University of Lavras for the experiment.

4.3. Greenhouse Experiment

We distributed the ramets among eighteen pots, which were each filled with 17 L of tap water. The water was treated by the local sanitation company and presented the following chemical composition: Cl—0.72 mg/L, F—0.74 ppm, and pH = 6.8. The ramets were left in the greenhouse for two weeks to allow an acclimation period without any interference. After this period, we selected 36 ramets of similar size, and isolated them in pots filled with 17 L of tap water. These ramets were cultivated until they produced asexual offspring, and were referred to as “parental” ramets from this point onward. The parental ramets remained attached to their offspring until a new generation of asexual offspring was produced.
Subsequently, both parental ramets and their offspring were isolated, forming basic experimental units. These groups of “mother-daughter” ramets were then placed in 36 pots, which were each filled with 17 L of tap water. We established four treatments for these parental-daughter ramet groups. The first treatment, termed “isolation treatment”, involved nine daughter ramets that were experimentally separated from their parental ramets. The second treatment, known as “clonal treatment”, maintained nine groups with the daughter ramets still attached to their parental ramets. The third treatment, referred to as “defoliation and isolation treatment”, consisted of nine daughter ramets that were both separated from their parental ramets and had all their leaves removed at the beginning of the experiment to simulate resource scarcity. Finally, the fourth treatment, called “defoliation treatment”, involved maintaining nine groups with the daughter ramets attached to their parental ramets, while also removing all leaves at the start of the experiment. We conducted the experiment over a duration of three months from 1 September to 30 November 2016. During this period of time, the mean temperature was 22 °C, and air humidity presented a mean of 66.5% [52].

4.4. Plant Biomass Measurement

At the end of the experiment, all plants were washed and transported to the laboratory. Each ramet was separated into its components: stem, petioles, leaf blades, and roots. These structures were placed in individual paper bags and dried at 60 °C for 60 h, or until a constant mass was achieved. Once dried, the mass of each ramet component was measured using a precision scale.
To assess the effects of our treatments on biomass allocation patterns, we calculated the proportion of biomass allocated to each structure for each individual ramet. This was accomplished by dividing the mass of each ramet component by the total biomass of the ramet, resulting in the proportion of biomass allocated to each measured part.

4.5. Data Analysis

All variables were previously tested for normality using Shapiro–Wilk tests. To test the influence of resource acquisition depletion on biomass allocation patterns in Eichhornia crassipes and its potential interactions with clonal integration, we employed generalized linear models (GLMs) with quasibinomial distribution for all variables. In each model, the biomass proportions allocated to each individual plant part were used as response variables, and an interaction term that considered the biomass proportions allocated to the other parts and treatments (isolated, clonal, isolated-defoliated, and clonal-defoliated) was included as the fixed variable. We then applied a stepwise backwards variable selection procedure for each model and simplified them to their most parsimonious form. All analyses were conducted using R Statistical Software (version 4.3.1) [53], and the graphs were built using the function ‘ggplot’ of the package ggplot2 [54].

5. Conclusions

We conclude that biomass allocation patterns to vegetative structures are mainly determined by resource pools. These relationships may respond in different ways to clonal integration, being less pronounced when the ramet is attached to its mother ramet, which indicates that clonal integration is an enhancer of E. crassipes growth and vegetative investment. This information sets a basis for aquatic plant management and control, and improves knowledge on clonal integration evolutionary importance and its significance to plant life strategies.

Author Contributions

Conceptualization: G.R.D. and F.d.F.C.; data curation: G.R.D.; formal analysis: G.R.D.; methodology: G.R.D. and F.d.F.C.; writing—original draft: G.R.D.; writing—review and editing: G.R.D., D.S. and F.d.F.C. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by Coordenação Nacional de Aperfeiçoamento de Pessoal de Nível Superior—Finance Code 001.

Data Availability Statement

The raw data supporting the conclusions of this article will be made available by the authors on request.

Acknowledgments

We thank CAPES for the grant given to the first author, Silvério José Coelho for logistic support for the greenhouse, Ana Fávaro and Eberton Borodinas for help with plant sampling, and Iara Ferreira for help during greenhouse experiments. We also thank Luziene Seixas for help with image editing. We also thank the three anonymous reviewers whose commentaries helped to improve this manuscript.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. (A) Relationship between the proportion of biomass allocated to the roots and leaf blades of E. crassipes ramets in relation to levels of clonal integration and experimental defoliation. (B) Relationship between the proportion of biomass allocated to the roots and petioles of E. crassipes ramets in relation to levels of clonal integration and experimental defoliation. (C) Relationship between the proportion of biomass allocated to the leaf blades and petioles of E. crassipes ramets in relation to levels of clonal integration and experimental defoliation. C represents the clonal treatment; I represents the isolated treatment; DI represents the defoliated and isolated treatment; and DC represents the defoliated treatment.
Figure 1. (A) Relationship between the proportion of biomass allocated to the roots and leaf blades of E. crassipes ramets in relation to levels of clonal integration and experimental defoliation. (B) Relationship between the proportion of biomass allocated to the roots and petioles of E. crassipes ramets in relation to levels of clonal integration and experimental defoliation. (C) Relationship between the proportion of biomass allocated to the leaf blades and petioles of E. crassipes ramets in relation to levels of clonal integration and experimental defoliation. C represents the clonal treatment; I represents the isolated treatment; DI represents the defoliated and isolated treatment; and DC represents the defoliated treatment.
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Table 1. Results of general linear models showing the relationships between the proportion of biomass allocated to each part of the plant. Significative relationships are marked with *.
Table 1. Results of general linear models showing the relationships between the proportion of biomass allocated to each part of the plant. Significative relationships are marked with *.
Y VariableX VariableTermEstimatet Valuep Value
Root biomass proportionStem biomass proportionIntercept0.60182.4720.0191 *
Stem biomass proportion−4.7915−1.7780.0853 *
Treatment (defoliated)−0.4651−2.7270.0104 *
Treatment (isolated and defoliated)0.26321.6490.1092
Treatment (isolated)0.11600.7790.4421
Leaf blade biomass proportionIntercept1.24773.95<0.001 *
Leaf blade biomass proportion−9.1477−3.438<0.01 *
Treatment (defoliated)−1.8855−4.29<0.001 *
Treatment (defoliated and isolated)−0.5727−1.2980.2065
Treatment (isolated)−0.3709−0.8030.4285
Leaf blade × Treatment (defoliated)11.34032.997<0.01 *
Leaf blade × Treatment (defoliated and isolated)4.08680.7250.474
Leaf blade × Treatment (isolated)3.56110.8490.403
Petioles biomass proportionIntercept1.76917.303<0.001 *
Petioles biomass proportion−7.0591−6.638<0.001 *
Treatment (defoliated)−0.8453−2.459<0.05 *
Treatment (defoliated and isolated)−0.2198−0.6370.592
Treatment (isolated)−0.2706−0.4780.63
Petioles × Treatment (defoliated)2.89532.221<0.05 *
Petioles × Treatment (defoliated and isolated)2.311.6120.1182
Petioles × Treatment (isolated)2.13890.90.3759
Leaf blade biomass proportionPetioles biomass proportionIntercept−3.3102−7.009<0.001 *
Petioles biomass proportion5.54782.766<0.01 *
Treatment (defoliated)2.28263.485<0.01 *
Treatment (defoliated and isolated)0.42070.5630.57817
Treatment (isolated)0.24580.2250.823
Petioles × Treatment (defoliated)−1.9822−3.647<0.01 *
Petioles × Treatment (defoliated and isolated)−9.0158−1.6580.108
Petioles × Treatment (isolated)−5.039−0.4360.666 *
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Demetrio, G.R.; Serafim, D.; Coelho, F.d.F. Is Clonal Integration a Buffer for the Stress of Resource Acquisition Depletion in Eichhornia crassipes (Pontederiaceae) Ramets? Stresses 2024, 4, 734-743. https://doi.org/10.3390/stresses4040047

AMA Style

Demetrio GR, Serafim D, Coelho FdF. Is Clonal Integration a Buffer for the Stress of Resource Acquisition Depletion in Eichhornia crassipes (Pontederiaceae) Ramets? Stresses. 2024; 4(4):734-743. https://doi.org/10.3390/stresses4040047

Chicago/Turabian Style

Demetrio, Guilherme Ramos, Dalton Serafim, and Flávia de Freitas Coelho. 2024. "Is Clonal Integration a Buffer for the Stress of Resource Acquisition Depletion in Eichhornia crassipes (Pontederiaceae) Ramets?" Stresses 4, no. 4: 734-743. https://doi.org/10.3390/stresses4040047

APA Style

Demetrio, G. R., Serafim, D., & Coelho, F. d. F. (2024). Is Clonal Integration a Buffer for the Stress of Resource Acquisition Depletion in Eichhornia crassipes (Pontederiaceae) Ramets? Stresses, 4(4), 734-743. https://doi.org/10.3390/stresses4040047

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