Cyanobacterial inoculation promotes enhanced clonal growth performance of the aquaticplant Salvinia auriculata.

Cyanobacterial inoculation promotes enhanced clonal growth performance of the aquaticplant Salvinia auriculata. | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Cyanobacterial inoculation promotes enhanced clonal growth performance of the aquaticplant Salvinia auriculata. Larissa Langsdorff Pimenta, Grécia Andrade Souza, Luis Carlos Pereira, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-5822876/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Clonal plants, like Salvinia auriculata , are widespread and perform important ecosystem functions, influencing the structure and composition of the ecosystems in which they occur. Some cyanobacteria perform biological nitrogen fixation (BNF) and can affect plant growth, as nitrogen (N) is a limiting nutrient. Therefore, we carried out a greenhouse experiment with the inoculation of the cyanobacterial strain Desmonostoc (UFLA 12) to investigate whether heterocystous cyanobacteria favour individual growth and drives reproductive strategies (sexual reproduction and/or clonal growth) of S. auriculata . Salvinia auriculata ramets were grown in plastic pots under the following treatments: (D) Desmonostoc (UFLA 12) inoculum and (Co) control, in which cyanobacteria were absent. The Desmonostoc presence positively influenced the clonal growth of S. auriculata , and increased shoots number, plants fresh biomass, and shoot size. We conclude that the inoculation of Desmonostoc (UFLA 12) contributes to a more vigorous spread of S. auriculata since it enhanced clonal growth. In this sense, the role of Desmonostoc (UFLA 12) as a potential biofertilizer may serve as a tool to assist in understanding the excessive growth of S. auriculata in aquatic environments. Aquatic Plant Biofertiliser Clonal Growth Heterocystous Cyanobacteria Reproductive Strategies Figures Figure 1 Figure 2 Introduction Aquatic macrophytes are widespread plants that inhabits water bodies in several different ecosystems (Alahuhta et al. 2021 ). Their distribution is linked to many factors that range from the landscape level, as altitude and water body area (Rolon and Maltchik 2006 ), to local factors as nutrient availability (Dar et al. 2014 ). Nutrient limitation seems to play a major role in most aquatic macrophyte species growth and development, with carbon, phosphorus and nitrogen being the most important ones (Bornette and Puijalon 2011 ). Nitrogen (N) is one of the most important nutrients for plant growth, required for the synthesis of essential cellular components such as nucleic acids (Kuypers et al. 2018 ). However, although it is widely available in the atmosphere in the form of dinitrogen (N 2 ), most living organisms cannot incorporate it into their metabolism because it is an inert gas (Hoffman et al. 2014 ). Biological nitrogen fixation (BNF), the conversion of N 2 into molecules that can be used by other organisms, such as ammonium ions and ammonia, is only performed by some prokaryotic microorganisms, including cyanobacteria (Mazhar et al. 2019 ). Cyanobacteria have the ability to form endophytic associations with various organisms, including plants (mosses, cycads, ferns), fungi (forming lichens), and algae (Adams 2000 ; Aguiar et al. 2008 ; Rai et al. 2002 ). In addition, several strains form epiphytic associations with floating aquatic plants, such as S. auriculata (Pimenta et al. 2022 ), and some even with submerged aquatic plants, such as Stratiotes aloides , showing allelopathic activity (Mohamed and Shehri 2010).Endophytic symbiotic associations of cyanobacteria are well studied as they contribute significantly to global BNF (Elbert et al. 2012 ; Kluge et al. 2002 ; Meeks 2005 ). Trichormus azollae ( Anabaena azollae ) is a heterocytous cyanobacterium that can reside in the leaf cavities of the aquatic fern Azolla , providing nitrogen for use as a biofertiliser in rice crops (Sergeeva et al. 2002 ; Ahmed et al. 2010 ; Kollah et al. 2016 ). In addition to nitrogen fixation, cyanobacteria secrete a variety of secondary metabolites, such as growth-promoting hormones, including auxins, gibberellic acid, and cytokinins (Haroun and Hussein 2003 ; Shariatmadari et al. 2013 ), benefiting the cultivation of various crops such as wheat (Kholssi et al. 2021 ), tomato (Prasanna et al. 2013 ), corn (Prasanna et al. 2016a ), chickpea (Prasanna et al. 2017 ), and cotton (Prasanna et al. 2016b ). The relationship between water nutrient content and aquatic macrophytes is also mediated by the plant’s life form, and free floating macrophytes depend upon a-high levels of nutrients in the water column to present high physiological performances (Lacoul and Freedman 2006 ). In addition to different life forms, aquatic macrophytes can also present different reproductive systems. In fact, the ability of clonal reproduction that is present in most of aquatic macrophytes is cited as one of the factors that allow the ecological success of these plants group (Santamaria 2002). In this sense, aquatic clonal plants are widely distributed and play essential ecosystem roles as they are key species in nutrient cycling and present high primary productivity, influencing the structure and composition of aquatic ecosystems (De Kroon and Hutchings 1995 ; Zuo et al. 2023 ). They reproduce vegetatively, producing genetically identical individuals named ramets (Zuo et al. 2023 ), which are physiologically independent of the mother plant (Franklin et al. 2021 ; Demetrio and Coelho 2023 ), or by sexual reproduction (Harper 1977 ; Demetrio and Coelho 2023 ). In environments with high nutrient availability, clonal plants may produce ramets as an adaptive strategy to utilise all available resources and become dominant over other species (Zheng et al. 2019 ; Zhang et al. 2020 ). Therefore, changes in environmental conditions, such as fertilisation (Gonzalez et al. 2016) and nutrient limitation (Dong et al. 1997 ), can affect plant clonal growth (Geng and He 2021 ). In this sense S. auriculata is a good model to understand the relationship between plants and nitrogen-fixing cyanobatcteria, as it is a common freshwater fern that, under favourable conditions, rapidly colonises large water surfaces through clonal growth (Medeiros et al. 2017 ). In addition, its submerged leaflets holds a periphyton that encompasses different species cyanobacteria that can influence its growth by providing fixed nitrogen (N) (Pimenta et al. 2022 ). Based on the premise that nitrogen-fixing cyanobacteria can influence plant growth (Ahmed et al. 2010 ; Diez and Ininbergs 2014; Rai et al. 2019 ), we hypothesized that the inoculation of Desmonostoc (UFLA 12) strain will enhance S auriculata individual performance and clonal growth, promoting an increase in both size and the number of new ramets. Materials and Methods Aquatic plant and their epiphytic cyanobacteria Salvinia auriculata Aubl. (Salviniaceae) is a free-floating aquatic fern (Coelho et al. 2005b ; Miranda and Schwartsburd 2019 ) that presents clonal reproduction by sprouting, in which each ramet can produce genetically identical ramets. It also presents sexual reproduction, in which spore-producing structures called sori are enclosed by a globular indusium, forming fertile fronds (De La Sota 1962 ; Miranda and Schwartsburd 2019 ). The ramets are connected by rhizomes that form colonies and consist of nodes, two aerial folioles responsible for photosynthesis, and submerged, finely divided folioles with a root function responsible for water and nutrient absorption (Room 1983 ; Sculthorpe 1967 ). We previously observed and isolated the Desmonostoc strain (UFLA 12) from the submerged folioles of S. auriculata , ( see in Pimenta et al. 2022 ). Desmonostoc (UFLA 12) strains have terminal and intercalary heterocysts, can perform BNF, and form long filaments surrounded by a diffusible mucilaginous sheath. Desmonostoc (UFLA 12) filaments are not densely coiled with compact trichomes as in Nostoc . In addition, akinetes can be differentiated in long chains as a form of resistance (Hrouzek et al. 2013 ). Cyanobacterial inoculum The cyanobacterial strain used in this study, Desmonostoc (UFLA 12) was previously isolated from the epiphyton of S. auriculata roots (Pimenta et al. 2022 ). The culture is deposited in the Collection of Cyanobacteria Cultures (CFC - UFLA), available at the Federal University of Lavras (UFLA). For cyanobacterial biomass production, the strain was inoculated by streaking on 100 Petri dishes (50 for each strain) containing solid BG-11 0 culture medium and placed on a bench to grow under a 12 h light/12 h dark photoperiod. After 60 days, the biomass was scraped from the plates and transferred to 2 litres of liquid BG-11 0 (1 litre for each strain) and homogenised using 20 mL syringes. To obtain an inoculum free of agar particles and dead cells, the pre-inoculum was centrifuged at 5000 rpm for 5 minutes (Thermo Scientific MULTIFUGE X1R centrifuge). After centrifugation, the supernatant was removed, and the pellet was resuspended in 5 L of liquid medium (2.5 L for each strain) to obtain the desired optical density, measured by light spectrophotometry at a wavelength of 680 nm. The cyanobacterial density of the inoculum was estimated using a spectrophotometer (SHIMADZU UV-1800), obtaining an optical density of 0.283. This inoculum was divided into 50 mL Falcon tubes containing 15 mL of inoculum (1% of the total volume of the trays) for each replicate of the greenhouse experiment. In total, there were 10 tubes containing 15 mL of BG-11 0 medium with Desmonostoc (UFLA 12), 10 tubes containing 7.5 mL of BG-11 0 medium with Desmonostoc (UFLA 12), and 10 tubes containing only 15 mL of BG-11 0 medium for the control treatment. Greenhouse experiment Salvinia auriculata ramets were collected in summer from a permanent lagoon in the south-eastern region of Brazil (21°08'56"S, 44°52'53"W). After collection, the plants were carefully washed with distilled water to remove solid particles, dead parts and any cyanobacteria and eukaryotic algae attached to the roots. Healthy and morphologically similar ramets were selected and weighed to ensure that each tray contained a colony of three ramets weighing between three and four grams. After this process, colonies were transferred to a greenhouse with a 30% shade condition, and were acclimatized during seven days, prior to the start of the experiment.. After the acclimatization phase, colonies containing three initial ramets, called mother plants, were placed in plastic trays containing 1.5 L of filtered tap water and subjected to two treatments: (D) − 15 mL of Desmonostoc (UFLA 12) inoculum, and (Co) - No cyanobacterial inoculum, only 15 mL of BG-11 0 . Each treatment had ten replicates, totalling 20 trays (N = 20). Water was added to the trays every seven days to maintain the 1.5 L level. At the end of the 28-day experiment, the number of new ramets (shoots) and sori was counted, and the length and width of the aerial and submerged folioles of both the mother plant and the shoots were measured using calipers. After the measurements, the plants were dried in an oven at 60°C for 48 hours and weighed. Individual growth (dry biomass and size of mother plant), clonal growth (number of shoots, dry biomass and size of shoots) and investment in sexual reproduction (number of sori) were evaluated. Clonal growth (mother plant plus + shoots) was calculated by subtracting the initial fresh weight from the final fresh weight and dividing by the total duration of the experiment (28 days), as shown in the equation below (Gufu et al. 2019 ). $$\:Tx=\frac{\:PFF-\:PFI}{t}$$ Data analysis All response variables (mother plant dry weight, mother plant aerial folioles length and width, mother plant submerged folioles length and width, number of shoots, shoots dry weight, shoots aerial folioles length and width, shoots submerged folioles length and width, sori number per colony, and growth rate) were checked for normality with Shapiro-Wilk tests. We built generalized linear models (GLM’s) to test the effect of treatments (predictor variables) on S. auriculata mother plants and shoot traits. We built an individual model for each response variable that was performed with a gaussian distribution when data were normal and with quasipoisson distribution when data was not normal, to account for overdispersion (Crawley, 2012 ). All GLM’s were performed in the R environment, using the glm function of the base package (R Core Team, 2023 ). After this, we used the function rsquared of the “piecewiseSEM” package (Lefcheck, 2016 ) to obtain the R² for the predictor variables (Treatment) in each model. Results Cyanobacteria did not promote the general individual growth of initial ramets (mother plants) of S. auriculata (Fig. 1 A-E), but enhanced the growth rate and increased the foliole widthtable (Table 1 ). The treatments did not affect dry biomass of the mother plant (F = 1.791 p = 0.1975, Fig. 1 B), aerial leaf length (F = 0.5066, p = 0.4857; Fig. 1 D),, submerged leaf width (F = 1.2639, p = 0.2757, Fig. 1 E), and submerged leaf lenght (F = 2.3186, p = 0.1452, Fig. 1 E). In contrast to the individual growth of the mother plant of S. auriculata , there were differences in the clonal growth of S. auriculata (Fig. 1 G). The number of shoots varied among treatments (Table 1 ), with the highest number of shoots observed in the presence of Desmonostoc (D treatment) compared to control treatment (Co treatment). Sexual reproduction, however, was not affected by the presence of Desmonostoc (F = 2.1687, p = 0.1581, Fig. 1 G). On the other hand, the presence of Desmonostoc was highly important for the growth of Salvinia auriculata clonal offspring. The shoots that grew in the presence of Desmonostoc presented larger aerial and submerged leaves length and width (Table 1 , Fig. 2 A-E). Discussion The presence of Desmonostoc presented different effects on Salvinia auriculata that depended upon the level of plant organization that was sampled. Salvinia auriculata mother ramets were not affected Desmonostoc presence, while the shoots generated by these ramets were greatly affected in a positive way by the Cianobatceria presence. According to Gadgil and Bossert ( 1970 ), the life history of an organism can be seen as the result of three biological processes: survival, growth, and reproduction, which compete for resources. Salvinia auriculata can differentially allocate resources between these biological processes (Medeiros et al. 2016 ), and since we set up the experiment with already developed and grown ramets capable of clonal and sexual reproduction (De La Sota 1962 ; Miranda and Schwartsburd 2019 ), these ramets were likely not growing because they were allocating more resources in clonal growth. In addition, investment in sori production tends to occur in dry conditions and at high population densities, as observed by Coelho and colleagues ( 2005b ). In habitats with high environmental variability, this strategy is crucial to the plant's persistence (Coelho et al., 2021 ). In our experiment, we did not induce drought and we did not observe excessive ramet growth, which explains the lack of differences in sori production between treatments. Clonal growth, resulting in the rapid spread of ramets, is an important feature of the life history of aquatic plants, allowing them to efficiently occupy space (Seastedt 2009 ). The inoculation of the Desmonostoc (UFLA 12) strain contributed to the clonal growth of S. auriculata in all aspects, including numerical increase, biomass, and shoot size, as well as growth rate. Under favourable conditions, S. auriculata exhibits vigorous clonal growth (Julien et al. 2002 ). In terms of nutrients, nitrogen (N) and phosphorus (P) are the most limiting factors affecting the growth of aquatic plants (Duan et al. 2007 ; Smith 2014 ). In addition to their ability to fix nitrogen and release it into the environment, cyanobacteria are also partially capable of converting insoluble mineral phosphorus into soluble forms that can be used by other organisms. (Cameron and Julian 1988 ; Rai et al. 2019 ; Yandigeri et al. 2011 ). Therefore, we suggest that aquatic plants may benefit from the FBN that occurs in the periphytic community formed by cyanobacteria (Srivastava et al. 2017 ). Hempel and colleagues ( 2008 ) reported positive effects of periphyton on aquatic plants, such as the provision of organic compounds and carbon dioxide, making nutrient cycling more efficient. Therefore, we suggest that the FBN performed by Desmonostoc (UFLA 12) improves the nutritional conditions of the environment and favours the clonal growth of S. auriculata . The strength of the plant-cyanobacteria interaction and the resulting effects depend on both the plant and the cyanobacteria and are often highly specific (Kollmen and Strieth 2020; Stewart et al. 1983 ). Initially, the most studied and used genera for fertilisation belonged only to the genera Nostoc and Anabaena . (Bergman and Rai 1993). However, other genera such as Cylindrospermum , Synechococcus , Aulosira , Scytonema , Tolypothrix , Westiellopsis , Calothrix , Phormidium , Oscillatoria , Plectonema , and Gloeotrichia have been used (Manjunath et al. 2001; Osman et al. 2010 ; Prasanna et al. 2016b ; Singha 2009 ; Verma et al. 2016 ). In this experiment, we used Desmonostoc (UFLA12), which is phylogenetically close to Nostoc . Nostoc is a genus with a wide geographical distribution and more than 250 described taxa; however, molecular analyses have shown that the genus is not homogeneous, and some new taxa belonging to Desmonostoc (Hrouzek et al. 2013 ) and Mojavia (Řeháková et al. 2007 ) are based on Nostoc species (Sant'Anna et al. 2020). We suggest that the increased clonal growth of S. auriculata was greater in the treatment with Desmonostoc inoculation due to the specificity of the plant-cyanobacteria relationship (Kollmen and Strieth 2022 ; Stewart et al. 1983 ). The Desmonostoc (UFLA 12) strain used in the experiment was isolated from the roots of this aquatic plant. In addition, Desmonostoc produces a mucilaginous sheath that can facilitate the attachment of cyanobacteria to surfaces and nutrient deprivation, and store them (Droop 1968 ; Kollmen and Strieth 2022 ; Schmitt and Flemming 1999 ; Schooling and Beveridge 2006 ). The Desmonostoc (UFLA 12) strain is nitrogen-fixing and therefore capable of performing FBN. It can produces both terminal and intercalary heterocytes (Hrouzek et al. 2013 ) what may explain the enhanced growth observed on the S. auriculata individuals submitted to the strain addition. Studies have shown that periphyton and aquatic plants compete for nutrients (O'Hare et al. 2018; Périllon and Hilt 2019 ; Romo et al. 2007; Xie et al. 2013 ). Thus, S. auriculata shoots may have invested available resources in root growth for nutrient uptake rather than in aerial folioles, as no different light conditions were imposed between treatments that could shape resource allocation for increased leaf area and photosynthetic rates (Medeiros et al. 2017 ). Biomass allocation to different parts of aquatic plants, such as roots and leaves, may occur due to limitation of available nutrients, as observed by Zhou and colleagues ( 2011 ). Several authors have reported plant growth in the presence of cyanobacteria. Soil fertilisation by different cyanobacterial strains increased root length, height, and fresh and dry biomass of pumpkin, cucumber, and tomato plants (Shariatmadari et al. 2013 ; Hashtroudi et al. 2013 ). Cylindrospermum muscicola and Anabaena oryzae increased the growth of Lupinus termis (Haroun and Hussein, 2003 ), and Nostoc entophytum and Oscillatoria angustissima contributed to the growth of peas. It was also observed that the consortium of Calotrix sp., Anabaena cylindrica , and rhizobacteria promoted the growth of wheat in a hydroponic system (Kholssi et al. 2021 ). Due to the social and environmental problems caused by the overuse of agricultural inputs, combined with the high cost of fertilisers, the search for more sustainable alternatives, such as biofertilisers, has increased, and the potential for fertilisation by cyanobacteria has been explored (Sadvakasova et al. 2023 ). We observed the fertilisation capacity of Desmonostoc (UFLA 12) in the growth of S. auriculata , contributing to its more vigorous clonal spread. The fact that Desmonostoc (UFLA 12) favoured the clonal growth of S. auriculata can be used as a tool to understand the excessive growth of these plants in aquatic environments, with the aim of biofertilising plants or managing them. In conclusion, the Desmonostoc (UFLA 12) strain proved to be an excellent growth promoter for S. auriculata , a wild plant. It is therefore a promising strain for use in fertilisation. Table 1 Table 1 Values ​​of the statistical tests, for the mother plant and shoots Ramet type Model Variation Source Estimate Std Error t-value p-value R 2 Mother plant Growth rate ~ Treatment Intercept 0.083 0.013 6.037 < 0.01 0.44 Desmonostoc presence 0.074 0.019 3.806 < 0.01 Mother plant aerial leaf width ~ Treatment Intercept 2.766 0.072 37.966 < 0.01 0.25 Desmonostoc presence -0.258 0.103 -2.504 < 0.05 Shoot Shoot dry weight ~Treatment Intercept -3.649 0.245 -14.857 < 0.01 0.23 Desmonostoc presence 0.785 0.296 2.648 < 0.01 Shoot aerial leaf width ~Treatment Intercept 0.306 0.294 1.039 0.312 0.57 Desmonostoc presence 1.072 0.341 3.145 < 0.01 Shoot aerial leaf length ~Treatment Intercept 0.124 0.248 0.498 0.624 0.45 Desmonostoc presence 0.911 0.294 3.09 < 0.01 Shoot submerse leaf width ~Treatment Intercept -1.435 0.298 -4.816 < 0.01 0.34 Desmonostoc presence 1.029 0.347 2.963 < 0.01 Shoot submerse leaf length ~Treatment Intercept -0.1803 0.306 -0.589 0.562 0.45 Desmonostoc presence 1.050 0.355 2.955 < 0.01 Declarations Competing Interests The authors declare no competing interests. Funding This work was supported by Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG – APQ-01347-22). Author Contribution LLPConceptualization, Methodology and Investigation (execution of the experiment) , Formalanalysis, Writing – original draft, ValidationGASMethodology and Investigation (Inoculum preparation and experiment maintenance)LCPMethodology and Investigation (Inoculum preparation and experiment maintenance)MGMVVMethodology (inoculum preparation) and Investigation (identification of Cronbergia sp.(UFLA 35) and Desmonostoc sp. (UFLA 12) strains.GRDConceptualization, Formal analysis, Writing – original draft, ValidationFFCConceptualization, Formal analysis, Writing – original draft, Validation Acknowledgement The authors are grateful to the professors Victor Satler Pylro, Joziana Muniz de Paiva Barcante and Whasley Ferreira Duarte, for providing the necessary research equipment; To the professors Flávia Maria Avelar Gonçalves and José Airton Rodrigues Nunes for providing green house. This work was supported by Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG – APQ-01347-22). Availability of data and material The data will be made available for consultation, if required Code availability Not applicable References Adams DG (2000) Symbiotic interactions. In: Whitton BA, Potts M (Eds.) The Ecology of Cyanobacteria, Their Diversity in Time and Space, Kluwer Academic Publishers, Dordrecht, the Netherlands, pp. 523–61. Aguiar R, Fiore MF, Franco MW, Ventrella MC, Lorenzi AS, Vanetti CA, Alfenas AC (2008) A Novel epiphytic cyanobacterial species from the genus Brasilonema causing damage to Eucalyptus leaves 1. Journal of Phycology, 44(5), 1322–1334. https://doi.org/10.1111/j.1529-8817.2008.00584.x Ahmed M, Stal LJ, Hasnain S (2010) Association of non-heterocystous cyanobacteria with crop plants. Plant and Soil, 336, 363–375. https://doi.org/10.1007/s11104-010-0488-x Alahuhta J, Lindholm M, Baastrup-Spohr L, García-Girón J, Toivanen M, Heino J, Murphy K (2021) Macroecology of macrophytes in the freshwater realm: Patterns, mechanisms and implications. Aquatic Botany, 168, 103325. Biondi M (2021) Adaptações de cianobactérias filamentosas e seus efeitos nas estratégias reprodutivas de Pistia stratiotes . 69 f. Tese (Doutorado) - Curso de Botânica Aplicada, Universidade Federal de Lavras. Bornette G, Puijalon S (2011) Response of aquatic plants to abiotic factors: a review. Aquatic sciences, 73, 1–14. https://doi.org/10.1007/s00027-010-0162-7 Cameron HJ, Julian GR (1988) Utilization of hydroxyapatite by cyanobacteria as their sole source of phosphate and calcium, Plant and soil, 109, 123–124. https://doi.org/10.1007/BF02197589 Coelho FF, Basulto SR, Fávaro A, Demetrio GR (2021) Sex allocation in aquatic ferns: The role of body size. Plant Species Biol. 2021; 36: 425–435. https://doi.org/10.1111/1442-1984.12324 Coelho FF, Deboni L, Santos Lopes F (2005a) Density-dependent reproductive and vegetative allocation in the aquatic plant Pistia stratiotes (Araceae), Revista de biología tropical, 53(3–4), 369–376. https://doi.org/10.15517/rbt.v53i3-4.14599 Coelho FF, Lopes FS, Sperber, CF (2005b) Persistence strategy of Salvinia auriculata Aublet in temporary ponds of Southern Pantanal, Brazil, Aquatic Botany, 81(4), 343–352. https://doi.org/10.1016/j.aquabot.2005.02.001 Crawley MJ (2012) The R book. John Wiley & Sons. Dar NA, Pandit AK, Ganai BA (2014) Factors Affecting the Distribution Patterns of Aquatic Macrophytes. Limnological Review, 14(2):75–81. https://doi.org/10.2478/limre-2014-0008 Demetrio GR, Coelho FF (2023) What are the consequences of clonal integration for floral traits and reproductive investment of a broadly distributed aquatic plant? Flora, 303, 152292 .https://doi.org/10.1016/j.flora.2023.152292 De Kroon H, Hutchings MJ (1995) Morphological plasticity in clonal plants: the foraging concept reconsidered, Journal of Ecology, 143–152. https://doi.org/10.2307/2261158 De La Sota ER (1962) Contribución al conocimiento de Las Salviniaceae neotropicales, I Salvinia oblongifolia Martius. Darwiniana, 465–498. Dıez B, Ininbergs K (2014) Ecological importance of cyanobacteria. In: Sharma NK, Rai AK, Stal LJ (eds) Cyanobacteria: an economic perspective. Wiley, Chichester. https://doi.org/10.1002/9781118402238.ch3 Dong M, During HJ, Werger MJ (1997) Clonal plasticity in response to nutrient availability in the pseudoannual herb, Trientalis europaea L, Plant Ecology, 131, 233–239. https://doi.org/10.1023/A:1009783921753 Droop MR (1968) Vitamin B12 and marine ecology. IV. The kinetics of uptake, growth and inhibition in Monochrysis lutheri , Journal of the Marine Biological Association of the United Kingdom, 48(3), 689–733. https://doi.org/10.1017/S0025315400019238 Duan YH, Zhang YL, Ye LT, Fan XR, Xu, GH, Shen, QR (2007). Responses of rice cultivars with different nitrogen use efficiency to partial nitrate nutrition, Annals of Botany, 99(6), 1153–1160. https://doi.org/10.1093/aob/mcm051 Elbert W, Weber B, Burrows S, Steinkamp J, Büdel B, Andreae MO, Pöschl U (2012) Contribution of cryptogamic covers to the global cycles of carbon and nitrogen, Nature Geoscience, 5(7), 459–462. https://doi.org/10.1038/ngeo1486 Franklin S, Alpert P, Salguero-Gómez R, Janovský Z, Herben T, Klimešová J, Douhovnikoff V (2021), Next-gen plant clonal ecology, Perspectives in plant ecology, evolution and systematics, 49, 125601. https://doi.org/10.1016/j.ppees.2021.125601 Gadgil M, Bossert WH (1970) Life historical consequences of natural selection, The American Naturalist, 104(935), 1–24. https://doi.org/10.1086/282637 Geng XM, He WM (2021) Success of native and invasive plant congeners depends on inorganic nitrogen compositions and levels, Journal of Plant Ecology, 14(2), 202–212. https://doi.org/10.1093/jpe/rtaa088 Genuário DB, Sant'Anna CL, Melo IS (2018) Elucidating the Cronbergia (Cyanobacteria) dilemma with the description of Cronbergia amazonensis sp. nov. isolated from Solimões river (Amazonia, Brazil), Algal research, 29, 233–241. https://doi.org/10.1016/j.algal.2017.11.034 González APR, Chrtek J, Dobrev, PI, Dumalasová V, Fehrer J, Mráz P, Latzel V (2016) Stress-induced memory alters growth of clonal offspring of white clover ( Trifolium repens ), American Journal of Botany, 103(9), 1567–1574. https://doi.org/10.3732/ajb.1500526 Gufu GD, Manea A, Leishman MR (2019) Growth, reproduction and functional trait responses of three freshwater plant species to elevated carbon dioxide, Aquatic Botany, 154, 18–23. https://doi.org/10.1016/j.aquabot.2018.12.006 Haroun SA, Hussein, MH (2003) The promotive effect of algal biofertilizers on growth, protein pattern and some metabolic activities of Lupinus termis plants grown in siliceous soil, Asian journal of Plant sciences. https://doi.org/10.3923/ajps.2003.944.951 Harper JL (1977) Population biology of plants, Population biology of plants. Hashtroudi MS, Ghassempour A, Riahi H, Shariatmadari Z, Khanjir M (2013) Endogenous auxins in plant growth-promoting Cyanobacteria— Anabaena vaginicola and Nostoc calcicola . Journal of applied phycology, 25, 379–386. https://doi.org/10.1007/s10811-012-9872-7 Hempel M, Blume M, Blindow I, Gross EM (2008) Epiphytic bacterial community composition on two common submerged macrophytes in brackish water and freshwater, Bmc Microbiology, 8(1), 1–10. https://doi.org/10.1186/1471-2180-8-58 Hoffman BM, Lukoyanov D, Yang ZY, Dean DR, Seefeldt LC (2014) Mechanism of nitrogen fixation by nitrogenase: the next stage. Chemical reviews, 114(8), 4041–4062. https://doi.org/10.1021/cr400641x Hrouzek P, Lukešová A, Mareš J, Ventura S (2013) Description of the cyanobacterial genus Desmonostoc gen. nov. including D. muscorum comb. nov. as a distinct, phylogenetically coherent taxon related to the genus Nostoc , Fottea, 13(2), 201–213. https://doi.org/10.5507/fot.2013.016 Julien MH, Center TD, Tipping PW (2002) Floating fern (salvinia), Biological control of invasive plants in the Eastern United States, 17–32. Kholssi R, Marks EA, Miñón J, Maté AP, Sacristán G, Montero O, Rad C (2021) A consortium of cyanobacteria and plant growth promoting rhizobacteria for wheat growth improvement in a hydroponic system, South African Journal of Botany, 142, 247–258. https://doi.org/10.1016/j.sajb.2021.06.035 Kluge M, Mollenhauer D, Wolf E, Schüßler A (2002) The Nostoc - Geosiphon endocytobiosis, IN: Cyanobacteria in symbiosis, 19–30. Kollah B, Patra AK, Mohanty SR (2016) Aquatic microphylla Azolla : a perspective paradigm for sustainable agriculture, environment and global climate change, Environmental Science and Pollution Research, 23, 4358–4369. https://doi.org/10.1007/s11356-015-5857-9 Kollmen J, Strieth D (2022) The beneficial effects of cyanobacterial co-culture on plant growth, Life, 12(2), 223. https://doi.org/10.3390/life12020223 Komárek J (2016) A polyphasic approach for the taxonomy of cyanobacteria: principles and applications, European Journal of Phycology, 51(3), 346–353. https://doi.org/10.1080/09670262.2016.1163738 Komárek J, Zapomelova E, Hindak F (2010) Cronbergia gen. nov., a new cyanobacterial genus (Cyanophyta) with a special strategy of heterocyte formation, Cryptogamie, Algologie, 31(3), 321–341. Komárek J, Johansen JR (2015) Filamentous cyanobacteria, In: Freshwater Algae of North America, Academic Press, 135–235. https://doi.org/10.1016/B978-0-12-385876-4.00004-9 Kuypers MM, Marchant HK, Kartal B (2018) The microbial nitrogen-cycling network, Nature Reviews Microbiology, 16(5), 263–276. https://doi.org/10.1038/nrmicro.2018.9 Lacoul P, Freedman B (2006) Relationships between aquatic plants and environmental factors along a steep Himalayan altitudinal gradient. Aquat Bot 84:3–16 Lefcheck JS (2016) “piecewiseSEM: Piecewise structural equation modeling in R for ecology, evolution, and systematics.” Methods in Ecology and Evolution , 7(5), 573–579. doi: 10.1111/2041-210X.12512 . Manjunath M, Prasanna R, Sharma P, Nain L, Singh R (2011) Developing PGPR consortia using novel genera Providencia and Alcaligenes along with cyanobacteria for wheat, Archives of Agronomy and Soil Science, 57(8), 873–887. https://doi.org/10.1080/03650340.2010.499902 Mazhar S, Cohen JD, Hasnain S (2019) Novel Approach for the Determination of Nitrogen Fixation in Cyanobacteria, Journal of the Chemical Society of Pakistan, 41(1), 105–105. Medeiros JCC, Coelho, FF, Teixeira E (2016) Biomass allocation and nutrients balance related to the concentration of Nitrogen and Phosphorus in Salvinia auriculata (Salviniaceae). Brazilian Journal of Biology, 76, 461–468. https://doi.org/10.1590/1519-6984.21114 Medeiros JCC, Silva JCF, Resende TDSC, Teodoro GS, Pereira FJ, Coelho, FF (2018) Ramet versus sporocarp production in the aquatic fern Salvinia auriculata (Salviniaceae): the role of shading, Australian Journal of Botany, 66(7), 583–588. https://doi.org/10.1071/BT18062 Medeiros JCC, Silva JCF, Teodoro GS, Coelho FF (2017) Effects of shade on individual ramet growth and on clonal growth of the aquatic fern Salvinia auriculata (Salviniaceae), American Fern Journal, 107(1), 21–29. https://doi.org/10.1640/0002-8444-107.1.21 Meeks JC (2005) Molecular mechanisms in the nitrogen-fixing Nostoc -bryophyte symbiosis, Molecular basis of symbiosis, 165–196. Meeks JC, Campbell EL, Summers ML, Wong FC (2002) Cellular differentiation in the cyanobacterium Nostoc punctiforme , Archives of microbiology, 178, 395–403. https://doi.org/10.1007/s00203-002-0476-5 Miranda CV, Schwartsburd PB (2019) Salvinia (Salviniaceae) in southern and southeastern Brazil—including new taxa, new distribution records, and new morphological characters, Brazilian Journal of Botany, 42, 171–188. https://doi.org/10.1007/s40415-019-00522-5 Mohamed ZA, Al Shehri AM (2010) Differential responses of epiphytic and planktonic toxic cyanobacteria to allelopathic substances of the submerged macrophyte Stratiotes aloides , International Review of Hydrobiology, 95(3), 224–234. https://doi.org/10.1002/iroh.200911219 O’Hare MT, Baattrup-Pedersen A, Baumgarte I, Freeman A, Gunn ID, Lázár AN, Bowes MJ (2018) Responses of aquatic plants to eutrophication in rivers: a revised conceptual model, Frontiers in plant science, 9, 451. https://doi.org/10.3389/fpls.2018.00451 Osman MEH, El-Sheekh MM, El-Naggar AH, Gheda SF (2010) Effect of two species of cyanobacteria as biofertilizers on some metabolic activities, growth, and yield of pea plant, Biology and fertility of soils, 46, 861–875. https://doi.org/10.1007/s00374-010-0491-7 Périllon C, Hilt S (2019) Groundwater discharge gives periphyton a competitive advantage over macrophytes, Aquatic Botany, 154, 72–80. https://doi.org/10.1016/j.aquabot.2019.01.004 Prasanna R, Babu S, Rana A, Kabi SR, Chaudhary V, Gupta V, Pal RK (2013). Evaluating the establishment and agronomic proficiency of cyanobacterial consortia as organic options in wheat–rice cropping sequence, Experimental agriculture, 49(3), 416–434. https://doi.org/10.1017/S001447971200107X Prasanna R, Kanchan A, Ramakrishnan B, Ranjan K, Venkatachalam S, Hossain F, Nain L (2016a) Cyanobacteria-based bioinoculants influence growth and yields by modulating the microbial communities favourably in the rhizospheres of maize hybrids, European Journal of Soil Biology, 75, 15–23. https://doi.org/10.1016/j.ejsobi.2016.04.001 Prasanna R, Ramakrishnan B, Ranjan K, Venkatachalam S, Kanchan A, Solanki P, Kranthi S (2016b) Microbial inoculants with multifaceted traits suppress Rhizoctonia populations and promote plant growth in cotton, Journal of Phytopathology, 164(11–12), 1030–1042. https://doi.org/10.1111/jph.12524 Prasanna R, Ramakrishnan B, Simranjit K, Ranjan K, Kanchan A, Hossain F, Nain L (2017) Cyanobacterial and rhizobial inoculation modulates the plant physiological attributes and nodule microbial communities of chickpea, Archives of Microbiology, 199, 1311–1323. https://doi.org/10.1007/s00203-017-1405-y Pimenta LL, Lima GP, Biondi M, Vaz MGMV, Coelho FF (2022) Epiphytic cyanobacterial strains in the roots of Salvinia auriculata and the effect of light and nutrients on the production of heterocyst, akinete and hormogonia, Aquatic Ecology, 56(3), 543–553. https://doi.org/10.1007/s10452-021-09922-w Pott VJ, Pott A (2000) Plantas aquáticas do Pantanal, Brasília, DF: Embrapa Comunicação para Transferência de Tecnologia; Corumbá: Embrapa-CPAP. Rai AN, Bergman B, Rasmussen U (2002) Cyanobacteria in symbiosis, Kluwer Academic Pub. Rai AN, Singh AK, Syiem MB (2019) Plant growth-promoting abilities in cyanobacteria, In: Cyanobacteria, Academic Press, 459–476 R Core Team (2023) R: A Language and Environment for Statistical Computing . R Foundation for Statistical Computing, Vienna, Austria. . Řeháková K, Johansen JR, Casamatta DA, Xuesong L, Vincent J (2007) Morphological and molecular characterization of selected desert soil cyanobacteria: three species new to science including Mojavia pulchra gen. et sp. nov.,Phycologia, 46(5), 481–502. https://doi.org/10.2216/06-92.1 Rolon AS, Maltchik L (2006) Environmental factors as predictors of aquatic macrophyte richness and composition in wetlands of southern Brazil. Hydrobiologia, 556, 221–231. Romo S (2007) Epiphyton, phytoplankton and macrophyte ecology in a shallow lake under in situ experimental conditions, Fundamental and applied limnology, 197–209. https://doi.org/10.1127/1863-9135/2007/0170-0197 Room PM (1983) Falling apart'as a lifestyle: the rhizome architecture and population growth of Salvinia molesta , The Journal of Ecology, 349–365. https://doi.org/10.2307/2259719 Sadvakasova AK, Bauenova MO, Kossalbayev BD, Zayadan BK, Huang Z, Wang J, Allakhverdiev SI (2023) Synthetic algocyanobacterial consortium as an alternative to chemical fertilizers, Environmental Research, 116418. https://doi.org/10.1016/j.envres.2023.116418 Sant’anna CL, Branco LHZ, Azevedo MTP, Werner VR, Malone CFS (2020) Cyanobacteria, In: Bicudo CEM, Menezes M, Gêneros de Algas de Águas Continentais no Brasil (4) p. 33–41. Santamaría L (2002) Why are most aquatic plants widely distributed? Dispersal, clonal growth and small-scale heterogeneity in a stressful environment. Acta oecologica, 23(3), 137–154. Schmitt J, Flemming HC (1999) Water binding in biofilms, Water science and technology, 39(7), 77–82. https://doi.org/10.1016/S0273-1223(99)00153-5 Schooling SR, Beveridge TJ (2006) Membrane vesicles: an overlooked component of the matrices of biofilms, Journal of bacteriology, 188(16), 5945–5957. https://doi.org/10.1128/jb.00257-06 Sculthorpe CD (1967) The biology of aquatic vascular plants. Seastedt T (2009) Traits of plant invaders, Nature, 459(7248), 783–784. https://doi.org/10.1038/459783a Sergeeva E, Liaimer A, Bergman B (2002) Evidence for production of the phytohormone indole-3-acetic acid by cyanobacteria, Planta, 215, 229–238. https://doi.org/10.1007/s00425-002-0749-x Shariatmadari Z, Riahi H, Seyed Hashtroudi M, Ghassempour A, Aghashariatmadary Z (2013) Plant growth promoting cyanobacteria and their distribution in terrestrial habitats of Iran, Soil Science and Plant Nutrition, 59(4), 535–547. https://doi.org/10.1080/00380768.2013.782253 Singha A (2009) Diversity of Cyanobacteria in Organic Farming Field Under Rice-wheat Cropping System (Doctoral dissertation, IARI, Division of Microbiology, New Delhi). Smith SD (2014) The roles of nitrogen and phosphorus in regulating the dominance of floating and submerged aquatic plants in a field mesocosm experiment, Aquatic Botany, 112, 1–9. https://doi.org/10.1016/j.aquabot.2013.07.001 Srivastava JK, Chandra H, Kalra SJ, Mishra P, Khan H, Yadav P (2017) Plant–microbe interaction in aquatic system and their role in the management of water quality: a review, Applied Water Science, 7, 1079–1090. https://doi.org/10.1007/s13201-016-0415-2 Stewart WDP, Rowell P, Rai AN (1983) Cyanobacteria-eukaryotic plant symbioses, In: Annales de l'Institut Pasteur/Microbiologie, Elsevier Masson 134 (1) 205–228. https://doi.org/10.1016/S0769-2609(83)80106-9 Verma S, Adak A, Prasanna R, Dhar S, Choudhary H, Nain L, Shivay YS (2016) Microbial priming elicits improved plant growth promotion and nutrient uptake in pea, Israel Journal of Plant Sciences, 63(3), 191–207. Wang R, Peng B, Huang K (2015) The research progress of CO2 sequestration by algal bio-fertilizer in China, Journal of CO2 Utilization, 11, 67–70. https://doi.org/10.1016/j.jcou.2015.01.007 Xie D, Yu D, You WH, Wang LG (2013) Algae mediate submerged macrophyte response to nutrient and dissolved inorganic carbon loading: a mesocosm study on different species, Chemosphere, 93(7), 1301–1308. https://doi.org/10.1016/j.chemosphere.2013.07.008 Yandigeri MS, Meena KK, Srinivasan R, Pabbi S (2011) Effect of mineral phosphate solubilization on biological nitrogen fixation by diazotrophic cyanobacteria, Indian journal of microbiology, 51, 48–53. https://doi.org/10.1007/s12088-011-0081-x Zhang LM, Jin Y, Yao SM, Lei NF, Chen JS, Zhang Q, Yu, FH (2020) Growth and morphological responses of duckweed to clonal fragmentation, nutrient availability, and population density, Frontiers in Plant Science, 11, 618. Zheng Z, Bai W, Zhang WH (2019) Clonality-dependent dynamic change of plant community in temperate grasslands under nitrogen enrichment, Oecologia, 189, 255–266. https://doi.org/10.1007/s00442-018-4317-x Zhou X, Wang G, Yang F (2011) Characteristics of growth, nutrient uptake, purification effect of Ipomoea aquatica , Lolium multiflorum , and Sorghum sudanense grown under different nitrogen levels, Desalination, 273(2–3), 366–374. https://doi.org/10.1016/j.desal.2011.01.057 Zuo S, Liu L, Zhou C, Wang X, Huang D (2023) Effects of nitrogen limitation on Leymus chinensis clonal growth form and bud bank, Flora, (298) 152186. https://doi.org/10.1016/j.flora.2022.152186 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-5822876","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":403009106,"identity":"3c78e7e9-5c0b-4a7b-96ca-03f8b474cea1","order_by":0,"name":"Larissa Langsdorff Pimenta","email":"","orcid":"","institution":"Universidade Federal de Lavras","correspondingAuthor":false,"prefix":"","firstName":"Larissa","middleName":"Langsdorff","lastName":"Pimenta","suffix":""},{"id":403009108,"identity":"a74f5242-aeba-4039-adc6-49ca6c105e57","order_by":1,"name":"Grécia Andrade Souza","email":"","orcid":"","institution":"Universidade Federal de Lavras","correspondingAuthor":false,"prefix":"","firstName":"Grécia","middleName":"Andrade","lastName":"Souza","suffix":""},{"id":403009110,"identity":"1ee82698-d7cb-4cb3-b6b9-ca5c76b569db","order_by":2,"name":"Luis Carlos Pereira","email":"","orcid":"","institution":"Universidade Federal de Lavras","correspondingAuthor":false,"prefix":"","firstName":"Luis","middleName":"Carlos","lastName":"Pereira","suffix":""},{"id":403009112,"identity":"c255cf34-0ae1-4096-8fb4-29c3266cdded","order_by":3,"name":"Marcelo Gomes Marçal Vieira Vaz","email":"","orcid":"","institution":"Biodiversita Tecnologia Microbiana","correspondingAuthor":false,"prefix":"","firstName":"Marcelo","middleName":"Gomes Marçal Vieira","lastName":"Vaz","suffix":""},{"id":403009114,"identity":"d75dc268-a13f-415c-ad43-5d696d4ae7ba","order_by":4,"name":"Guilherme Ramos Demetrio","email":"","orcid":"","institution":"Universidade Federal de Alagoas","correspondingAuthor":false,"prefix":"","firstName":"Guilherme","middleName":"Ramos","lastName":"Demetrio","suffix":""},{"id":403009116,"identity":"2817d54d-9c9c-4aeb-a4ba-211027874321","order_by":5,"name":"Flávia Freitas Coelho","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA5klEQVRIie3QsQrCMBCA4SuFZKlmTRH0FRIEFwdfJcXB2c1BRJd0EWcF8RmcnJVCJx/AzRTBWRHEQcRTcBFJdXPIz3Hbx5EAuFx/GI1xKSjj9jVAB1gRgFhJkDxJFbeHZA2h/orAi+B8QXx/z01XVBgdaHOcJZzQJdl1rITUuEqFnAxXsZwskASKyrWFNHyoCkWuSmwiXSoskh4BRcK+9Qo9CXUTSmwzJFO8wkweCaSJNJKNh6SPhOdeCdomGj3eEulwnLaQZLG0EhrPV5fz48eSlB+6dc5YM81s5FPer8Dlcrlc790BeQNEsOg78VwAAAAASUVORK5CYII=","orcid":"","institution":"Universidade Federal de Lavras","correspondingAuthor":true,"prefix":"","firstName":"Flávia","middleName":"Freitas","lastName":"Coelho","suffix":""}],"badges":[],"createdAt":"2025-01-13 23:08:06","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-5822876/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-5822876/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":74088487,"identity":"e6f5b8bf-87e8-476c-a32d-020ac50a9085","added_by":"auto","created_at":"2025-01-17 15:40:52","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":276136,"visible":true,"origin":"","legend":"\u003cp\u003eIndividual growth of the mother plant of \u003cem\u003eS. auriculata\u003c/em\u003e in each treatment. (A)Mean growth rate (cm/day); (B) Mother plant dry weight (g); (C) Mother plant aerial leaf width (cm); (D) Mother plant aerial leaf length (cm); (E) Mother plant submerse leaf width (cm); (F) Mother plant submerse leaf length (cm); (G) Number of shoots/Mother plant; (H) Number of sori/Mother plant\u003c/p\u003e","description":"","filename":"Fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-5822876/v1/c8d47320cfce2f51be5fb83c.png"},{"id":74088492,"identity":"61356872-c95b-4811-9b83-2772ea2b0a76","added_by":"auto","created_at":"2025-01-17 15:40:52","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":187029,"visible":true,"origin":"","legend":"\u003cp\u003eClonal growth of \u003cem\u003eS. auriculata\u003c/em\u003e in each treatment. (A) Shoot dry weight (g); (B) Shoot aerial leaf width (cm); (C) Shoot aerial leaf length (cm); (D) Shoot submerse leaf width (cm); (E) Shoot submerse leaf length (cm)\u003c/p\u003e","description":"","filename":"Fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-5822876/v1/4f5a1328afc3b6800bf9f1b5.png"},{"id":76013199,"identity":"8e29988b-103e-4063-8902-5ee4972bc6f3","added_by":"auto","created_at":"2025-02-11 12:31:53","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1135677,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5822876/v1/588a7cf4-a274-4234-9609-d0e69321b577.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Cyanobacterial inoculation promotes enhanced clonal growth performance of the aquaticplant Salvinia auriculata.","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAquatic macrophytes are widespread plants that inhabits water bodies in several different ecosystems (Alahuhta et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Their distribution is linked to many factors that range from the landscape level, as altitude and water body area (Rolon and Maltchik \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), to local factors as nutrient availability (Dar et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Nutrient limitation seems to play a major role in most aquatic macrophyte species growth and development, with carbon, phosphorus and nitrogen being the most important ones (Bornette and Puijalon \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Nitrogen (N) is one of the most important nutrients for plant growth, required for the synthesis of essential cellular components such as nucleic acids (Kuypers et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, although it is widely available in the atmosphere in the form of dinitrogen (N\u003csub\u003e2\u003c/sub\u003e), most living organisms cannot incorporate it into their metabolism because it is an inert gas (Hoffman et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Biological nitrogen fixation (BNF), the conversion of N\u003csub\u003e2\u003c/sub\u003e into molecules that can be used by other organisms, such as ammonium ions and ammonia, is only performed by some prokaryotic microorganisms, including cyanobacteria (Mazhar et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Cyanobacteria have the ability to form endophytic associations with various organisms, including plants (mosses, cycads, ferns), fungi (forming lichens), and algae (Adams \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Aguiar et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Rai et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). In addition, several strains form epiphytic associations with floating aquatic plants, such as \u003cem\u003eS. auriculata\u003c/em\u003e (Pimenta et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), and some even with submerged aquatic plants, such as \u003cem\u003eStratiotes aloides\u003c/em\u003e, showing allelopathic activity (Mohamed and Shehri 2010).Endophytic symbiotic associations of cyanobacteria are well studied as they contribute significantly to global BNF (Elbert et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Kluge et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Meeks \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). \u003cem\u003eTrichormus azollae\u003c/em\u003e (\u003cem\u003eAnabaena azollae\u003c/em\u003e) is a heterocytous cyanobacterium that can reside in the leaf cavities of the aquatic fern \u003cem\u003eAzolla\u003c/em\u003e, providing nitrogen for use as a biofertiliser in rice crops (Sergeeva et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Ahmed et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Kollah et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). In addition to nitrogen fixation, cyanobacteria secrete a variety of secondary metabolites, such as growth-promoting hormones, including auxins, gibberellic acid, and cytokinins (Haroun and Hussein \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Shariatmadari et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), benefiting the cultivation of various crops such as wheat (Kholssi et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), tomato (Prasanna et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), corn (Prasanna et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e), chickpea (Prasanna et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and cotton (Prasanna et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2016b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe relationship between water nutrient content and aquatic macrophytes is also mediated by the plant\u0026rsquo;s life form, and free floating macrophytes depend upon a-high levels of nutrients in the water column to present high physiological performances (Lacoul and Freedman \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). In addition to different life forms, aquatic macrophytes can also present different reproductive systems. In fact, the ability of clonal reproduction that is present in most of aquatic macrophytes is cited as one of the factors that allow the ecological success of these plants group (Santamaria 2002). In this sense, aquatic clonal plants are widely distributed and play essential ecosystem roles as they are key species in nutrient cycling and present high primary productivity, influencing the structure and composition of aquatic ecosystems (De Kroon and Hutchings \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Zuo et al. \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). They reproduce vegetatively, producing genetically identical individuals named ramets (Zuo et al. \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), which are physiologically independent of the mother plant (Franklin et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Demetrio and Coelho \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), or by sexual reproduction (Harper \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1977\u003c/span\u003e; Demetrio and Coelho \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In environments with high nutrient availability, clonal plants may produce ramets as an adaptive strategy to utilise all available resources and become dominant over other species (Zheng et al. \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Therefore, changes in environmental conditions, such as fertilisation (Gonzalez et al. 2016) and nutrient limitation (Dong et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), can affect plant clonal growth (Geng and He \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In this sense \u003cem\u003eS. auriculata\u003c/em\u003e is a good model to understand the relationship between plants and nitrogen-fixing cyanobatcteria, as it is a common freshwater fern that, under favourable conditions, rapidly colonises large water surfaces through clonal growth (Medeiros et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). In addition, its submerged leaflets holds a periphyton that encompasses different species cyanobacteria that can influence its growth by providing fixed nitrogen (N) (Pimenta et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBased on the premise that nitrogen-fixing cyanobacteria can influence plant growth (Ahmed et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Diez and Ininbergs 2014; Rai et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), we hypothesized that the inoculation of \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) strain will enhance \u003cem\u003eS auriculata\u003c/em\u003e individual performance and clonal growth, promoting an increase in both size and the number of new ramets.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eAquatic plant and their epiphytic cyanobacteria\u003c/h2\u003e \u003cp\u003e \u003cem\u003eSalvinia auriculata\u003c/em\u003e Aubl. (Salviniaceae) is a free-floating aquatic fern (Coelho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2005b\u003c/span\u003e; Miranda and Schwartsburd \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) that presents clonal reproduction by sprouting, in which each ramet can produce genetically identical ramets. It also presents sexual reproduction, in which spore-producing structures called sori are enclosed by a globular indusium, forming fertile fronds (De La Sota \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1962\u003c/span\u003e; Miranda and Schwartsburd \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The ramets are connected by rhizomes that form colonies and consist of nodes, two aerial folioles responsible for photosynthesis, and submerged, finely divided folioles with a root function responsible for water and nutrient absorption (Room \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Sculthorpe \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e1967\u003c/span\u003e). We previously observed and isolated the \u003cem\u003eDesmonostoc\u003c/em\u003e strain (UFLA 12) from the submerged folioles of \u003cem\u003eS. auriculata\u003c/em\u003e, (\u003cem\u003esee in\u003c/em\u003e Pimenta et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) strains have terminal and intercalary heterocysts, can perform BNF, and form long filaments surrounded by a diffusible mucilaginous sheath. \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) filaments are not densely coiled with compact trichomes as in \u003cem\u003eNostoc\u003c/em\u003e. In addition, akinetes can be differentiated in long chains as a form of resistance (Hrouzek et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eCyanobacterial inoculum\u003c/h3\u003e\n\u003cp\u003eThe cyanobacterial strain used in this study, \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) was previously isolated from the epiphyton of \u003cem\u003eS. auriculata\u003c/em\u003e roots (Pimenta et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The culture is deposited in the Collection of Cyanobacteria Cultures (CFC - UFLA), available at the Federal University of Lavras (UFLA). For cyanobacterial biomass production, the strain was inoculated by streaking on 100 Petri dishes (50 for each strain) containing solid BG-11\u003csub\u003e0\u003c/sub\u003e culture medium and placed on a bench to grow under a 12 h light/12 h dark photoperiod. After 60 days, the biomass was scraped from the plates and transferred to 2 litres of liquid BG-11\u003csub\u003e0\u003c/sub\u003e (1 litre for each strain) and homogenised using 20 mL syringes. To obtain an inoculum free of agar particles and dead cells, the pre-inoculum was centrifuged at 5000 rpm for 5 minutes (Thermo Scientific MULTIFUGE X1R centrifuge). After centrifugation, the supernatant was removed, and the pellet was resuspended in 5 L of liquid medium (2.5 L for each strain) to obtain the desired optical density, measured by light spectrophotometry at a wavelength of 680 nm. The cyanobacterial density of the inoculum was estimated using a spectrophotometer (SHIMADZU UV-1800), obtaining an optical density of 0.283. This inoculum was divided into 50 mL Falcon tubes containing 15 mL of inoculum (1% of the total volume of the trays) for each replicate of the greenhouse experiment. In total, there were 10 tubes containing 15 mL of BG-11\u003csub\u003e0\u003c/sub\u003e medium with \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12), 10 tubes containing 7.5 mL of BG-11\u003csub\u003e0\u003c/sub\u003e medium with \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12), and 10 tubes containing only 15 mL of BG-11\u003csub\u003e0\u003c/sub\u003e medium for the control treatment.\u003c/p\u003e\n\u003ch3\u003eGreenhouse experiment\u003c/h3\u003e\n\u003cp\u003e \u003cem\u003eSalvinia auriculata\u003c/em\u003e ramets were collected in summer from a permanent lagoon in the south-eastern region of Brazil (21\u0026deg;08'56\"S, 44\u0026deg;52'53\"W). After collection, the plants were carefully washed with distilled water to remove solid particles, dead parts and any cyanobacteria and eukaryotic algae attached to the roots. Healthy and morphologically similar ramets were selected and weighed to ensure that each tray contained a colony of three ramets weighing between three and four grams. After this process, colonies were transferred to a greenhouse with a 30% shade condition, and were acclimatized during seven days, prior to the start of the experiment..\u003c/p\u003e \u003cp\u003eAfter the acclimatization phase, colonies containing three initial ramets, called mother plants, were placed in plastic trays containing 1.5 L of filtered tap water and subjected to two treatments: (D) \u0026minus;\u0026thinsp;15 mL of \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) inoculum, and (Co) - No cyanobacterial inoculum, only 15 mL of BG-11\u003csub\u003e0\u003c/sub\u003e. Each treatment had ten replicates, totalling 20 trays (N\u0026thinsp;=\u0026thinsp;20). Water was added to the trays every seven days to maintain the 1.5 L level. At the end of the 28-day experiment, the number of new ramets (shoots) and sori was counted, and the length and width of the aerial and submerged folioles of both the mother plant and the shoots were measured using calipers. After the measurements, the plants were dried in an oven at 60\u0026deg;C for 48 hours and weighed. Individual growth (dry biomass and size of mother plant), clonal growth (number of shoots, dry biomass and size of shoots) and investment in sexual reproduction (number of sori) were evaluated. Clonal growth (mother plant plus\u0026thinsp;+\u0026thinsp;shoots) was calculated by subtracting the initial fresh weight from the final fresh weight and dividing by the total duration of the experiment (28 days), as shown in the equation below (Gufu et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003cdiv id=\"Equa\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equa\" name=\"EquationSource\"\u003e\n$$\\:Tx=\\frac{\\:PFF-\\:PFI}{t}$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eAll response variables (mother plant dry weight, mother plant aerial folioles length and width, mother plant submerged folioles length and width, number of shoots, shoots dry weight, shoots aerial folioles length and width, shoots submerged folioles length and width, sori number per colony, and growth rate) were checked for normality with Shapiro-Wilk tests. We built generalized linear models (GLM\u0026rsquo;s) to test the effect of treatments (predictor variables) on \u003cem\u003eS. auriculata\u003c/em\u003e mother plants and shoot traits. We built an individual model for each response variable that was performed with a gaussian distribution when data were normal and with quasipoisson distribution when data was not normal, to account for overdispersion (Crawley, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). All GLM\u0026rsquo;s were performed in the R environment, using the \u003cem\u003eglm\u003c/em\u003e function of the base package (R Core Team, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). After this, we used the function \u003cem\u003ersquared\u003c/em\u003e of the \u0026ldquo;piecewiseSEM\u0026rdquo; package (Lefcheck, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) to obtain the R\u0026sup2; for the predictor variables (Treatment) in each model.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eCyanobacteria did not promote the general individual growth of initial ramets (mother plants) of \u003cem\u003eS. auriculata\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA-E), but enhanced the growth rate and increased the foliole widthtable (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The treatments did not affect dry biomass of the mother plant (F\u0026thinsp;=\u0026thinsp;1.791 p\u0026thinsp;=\u0026thinsp;0.1975, Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB), aerial leaf length (F\u0026thinsp;=\u0026thinsp;0.5066, p\u0026thinsp;=\u0026thinsp;0.4857; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eD),, submerged leaf width (F\u0026thinsp;=\u0026thinsp;1.2639, p\u0026thinsp;=\u0026thinsp;0.2757, Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eE), and submerged leaf lenght (F\u0026thinsp;=\u0026thinsp;2.3186, p\u0026thinsp;=\u0026thinsp;0.1452, Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eE). In contrast to the individual growth of the mother plant of \u003cem\u003eS. auriculata\u003c/em\u003e, there were differences in the clonal growth of \u003cem\u003eS. auriculata\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eG). The number of shoots varied among treatments (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), with the highest number of shoots observed in the presence of \u003cem\u003eDesmonostoc\u003c/em\u003e (D treatment) compared to control treatment (Co treatment). Sexual reproduction, however, was not affected by the presence of \u003cem\u003eDesmonostoc\u003c/em\u003e (F\u0026thinsp;=\u0026thinsp;2.1687, p\u0026thinsp;=\u0026thinsp;0.1581, Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eG). On the other hand, the presence of \u003cem\u003eDesmonostoc\u003c/em\u003e was highly important for the growth of \u003cem\u003eSalvinia auriculata\u003c/em\u003e clonal offspring. The shoots that grew in the presence of \u003cem\u003eDesmonostoc\u003c/em\u003e presented larger aerial and submerged leaves length and width (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA-E).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe presence of \u003cem\u003eDesmonostoc\u003c/em\u003e presented different effects on \u003cem\u003eSalvinia auriculata\u003c/em\u003e that depended upon the level of plant organization that was sampled. \u003cem\u003eSalvinia auriculata\u003c/em\u003e mother ramets were not affected \u003cem\u003eDesmonostoc\u003c/em\u003e presence, while the shoots generated by these ramets were greatly affected in a positive way by the Cianobatceria presence. According to Gadgil and Bossert (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1970\u003c/span\u003e), the life history of an organism can be seen as the result of three biological processes: survival, growth, and reproduction, which compete for resources. \u003cem\u003eSalvinia auriculata\u003c/em\u003e can differentially allocate resources between these biological processes (Medeiros et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), and since we set up the experiment with already developed and grown ramets capable of clonal and sexual reproduction (De La Sota \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1962\u003c/span\u003e; Miranda and Schwartsburd \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), these ramets were likely not growing because they were allocating more resources in clonal growth. In addition, investment in sori production tends to occur in dry conditions and at high population densities, as observed by Coelho and colleagues (\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2005b\u003c/span\u003e). In habitats with high environmental variability, this strategy is crucial to the plant's persistence (Coelho et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In our experiment, we did not induce drought and we did not observe excessive ramet growth, which explains the lack of differences in sori production between treatments.\u003c/p\u003e \u003cp\u003eClonal growth, resulting in the rapid spread of ramets, is an important feature of the life history of aquatic plants, allowing them to efficiently occupy space (Seastedt \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). The inoculation of the \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) strain contributed to the clonal growth of \u003cem\u003eS. auriculata\u003c/em\u003e in all aspects, including numerical increase, biomass, and shoot size, as well as growth rate. Under favourable conditions, \u003cem\u003eS. auriculata\u003c/em\u003e exhibits vigorous clonal growth (Julien et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). In terms of nutrients, nitrogen (N) and phosphorus (P) are the most limiting factors affecting the growth of aquatic plants (Duan et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Smith \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). In addition to their ability to fix nitrogen and release it into the environment, cyanobacteria are also partially capable of converting insoluble mineral phosphorus into soluble forms that can be used by other organisms. (Cameron and Julian \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Rai et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Yandigeri et al. \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Therefore, we suggest that aquatic plants may benefit from the FBN that occurs in the periphytic community formed by cyanobacteria (Srivastava et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Hempel and colleagues (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) reported positive effects of periphyton on aquatic plants, such as the provision of organic compounds and carbon dioxide, making nutrient cycling more efficient. Therefore, we suggest that the FBN performed by \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) improves the nutritional conditions of the environment and favours the clonal growth of \u003cem\u003eS. auriculata\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThe strength of the plant-cyanobacteria interaction and the resulting effects depend on both the plant and the cyanobacteria and are often highly specific (Kollmen and Strieth 2020; Stewart et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e1983\u003c/span\u003e). Initially, the most studied and used genera for fertilisation belonged only to the genera \u003cem\u003eNostoc\u003c/em\u003e and \u003cem\u003eAnabaena\u003c/em\u003e. (Bergman and Rai 1993). However, other genera such as \u003cem\u003eCylindrospermum\u003c/em\u003e, \u003cem\u003eSynechococcus\u003c/em\u003e, \u003cem\u003eAulosira\u003c/em\u003e, \u003cem\u003eScytonema\u003c/em\u003e, \u003cem\u003eTolypothrix\u003c/em\u003e, \u003cem\u003eWestiellopsis\u003c/em\u003e, \u003cem\u003eCalothrix\u003c/em\u003e, \u003cem\u003ePhormidium\u003c/em\u003e, \u003cem\u003eOscillatoria\u003c/em\u003e, \u003cem\u003ePlectonema\u003c/em\u003e, and \u003cem\u003eGloeotrichia\u003c/em\u003e have been used (Manjunath et al. 2001; Osman et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Prasanna et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2016b\u003c/span\u003e; Singha \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Verma et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn this experiment, we used \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA12), which is phylogenetically close to \u003cem\u003eNostoc\u003c/em\u003e. \u003cem\u003eNostoc\u003c/em\u003e is a genus with a wide geographical distribution and more than 250 described taxa; however, molecular analyses have shown that the genus is not homogeneous, and some new taxa belonging to \u003cem\u003eDesmonostoc\u003c/em\u003e (Hrouzek et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and \u003cem\u003eMojavia\u003c/em\u003e (Řeh\u0026aacute;kov\u0026aacute; et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) are based on \u003cem\u003eNostoc\u003c/em\u003e species (Sant'Anna et al. 2020).\u003c/p\u003e \u003cp\u003eWe suggest that the increased clonal growth of \u003cem\u003eS. auriculata\u003c/em\u003e was greater in the treatment with \u003cem\u003eDesmonostoc\u003c/em\u003e inoculation due to the specificity of the plant-cyanobacteria relationship (Kollmen and Strieth \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Stewart et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e1983\u003c/span\u003e). The \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) strain used in the experiment was isolated from the roots of this aquatic plant. In addition, \u003cem\u003eDesmonostoc\u003c/em\u003e produces a mucilaginous sheath that can facilitate the attachment of cyanobacteria to surfaces and nutrient deprivation, and store them (Droop \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1968\u003c/span\u003e; Kollmen and Strieth \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Schmitt and Flemming \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Schooling and Beveridge \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) strain is nitrogen-fixing and therefore capable of performing FBN. It can produces both terminal and intercalary heterocytes (Hrouzek et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) what may explain the enhanced growth observed on the \u003cem\u003eS. auriculata\u003c/em\u003e individuals submitted to the strain addition. Studies have shown that periphyton and aquatic plants compete for nutrients (O'Hare et al. 2018; P\u0026eacute;rillon and Hilt \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Romo et al. 2007; Xie et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Thus, \u003cem\u003eS. auriculata\u003c/em\u003e shoots may have invested available resources in root growth for nutrient uptake rather than in aerial folioles, as no different light conditions were imposed between treatments that could shape resource allocation for increased leaf area and photosynthetic rates (Medeiros et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Biomass allocation to different parts of aquatic plants, such as roots and leaves, may occur due to limitation of available nutrients, as observed by Zhou and colleagues (\u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeveral authors have reported plant growth in the presence of cyanobacteria. Soil fertilisation by different cyanobacterial strains increased root length, height, and fresh and dry biomass of pumpkin, cucumber, and tomato plants (Shariatmadari et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Hashtroudi et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). \u003cem\u003eCylindrospermum muscicola\u003c/em\u003e and \u003cem\u003eAnabaena oryzae\u003c/em\u003e increased the growth of \u003cem\u003eLupinus termis\u003c/em\u003e (Haroun and Hussein, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), and \u003cem\u003eNostoc entophytum\u003c/em\u003e and \u003cem\u003eOscillatoria angustissima\u003c/em\u003e contributed to the growth of peas. It was also observed that the consortium of \u003cem\u003eCalotrix\u003c/em\u003e sp., \u003cem\u003eAnabaena cylindrica\u003c/em\u003e, and rhizobacteria promoted the growth of wheat in a hydroponic system (Kholssi et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Due to the social and environmental problems caused by the overuse of agricultural inputs, combined with the high cost of fertilisers, the search for more sustainable alternatives, such as biofertilisers, has increased, and the potential for fertilisation by cyanobacteria has been explored (Sadvakasova et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWe observed the fertilisation capacity of \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) in the growth of \u003cem\u003eS. auriculata\u003c/em\u003e, contributing to its more vigorous clonal spread. The fact that \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) favoured the clonal growth of \u003cem\u003eS. auriculata\u003c/em\u003e can be used as a tool to understand the excessive growth of these plants in aquatic environments, with the aim of biofertilising plants or managing them.\u003c/p\u003e \u003cp\u003eIn conclusion, the \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) strain proved to be an excellent growth promoter for \u003cem\u003eS. auriculata\u003c/em\u003e, a wild plant. It is therefore a promising strain for use in fertilisation.\u003c/p\u003e \u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eValues ​​of the statistical tests, for the mother plant and shoots\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRamet type\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eModel\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eVariation Source\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEstimate\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eStd Error\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003et-value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003ep-value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eR\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003eMother plant\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGrowth rate\u0026thinsp;~\u0026thinsp;Treatment\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.083\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.013\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e6.037\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0.44\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDesmonostoc presence\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.074\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.019\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3.806\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMother plant aerial leaf width\u0026thinsp;~\u0026thinsp;Treatment\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2.766\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.072\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e37.966\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0.25\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDesmonostoc presence\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-0.258\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.103\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-2.504\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.05\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"9\" rowspan=\"10\"\u003e \u003cp\u003eShoot\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eShoot dry weight ~Treatment\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-3.649\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.245\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-14.857\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0.23\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDesmonostoc presence\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.785\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.296\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2.648\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eShoot aerial leaf width ~Treatment\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.306\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.294\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1.039\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.312\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0.57\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDesmonostoc presence\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.072\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.341\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3.145\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eShoot aerial leaf length ~Treatment\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.124\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.248\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.498\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.624\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0.45\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDesmonostoc presence\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.911\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.294\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eShoot submerse leaf width ~Treatment\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-1.435\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.298\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-4.816\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0.34\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDesmonostoc presence\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.029\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.347\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2.963\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eShoot submerse leaf length ~Treatment\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-0.1803\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.306\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-0.589\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.562\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0.45\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDesmonostoc presence\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.050\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.355\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2.955\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.01\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting Interests\u003c/h2\u003e \u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis work was supported by Funda\u0026ccedil;\u0026atilde;o de Amparo \u0026agrave; Pesquisa do Estado de Minas Gerais (FAPEMIG \u0026ndash; APQ-01347-22).\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eLLPConceptualization, Methodology and Investigation (execution of the experiment) , Formalanalysis, Writing \u0026ndash; original draft, ValidationGASMethodology and Investigation (Inoculum preparation and experiment maintenance)LCPMethodology and Investigation (Inoculum preparation and experiment maintenance)MGMVVMethodology (inoculum preparation) and Investigation (identification of Cronbergia sp.(UFLA 35) and Desmonostoc sp. (UFLA 12) strains.GRDConceptualization, Formal analysis, Writing \u0026ndash; original draft, ValidationFFCConceptualization, Formal analysis, Writing \u0026ndash; original draft, Validation\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eThe authors are grateful to the professors Victor Satler Pylro, Joziana Muniz de Paiva Barcante and Whasley Ferreira Duarte, for providing the necessary research equipment; To the professors Fl\u0026aacute;via Maria Avelar Gon\u0026ccedil;alves and Jos\u0026eacute; Airton Rodrigues Nunes for providing green house. This work was supported by Funda\u0026ccedil;\u0026atilde;o de Amparo \u0026agrave; Pesquisa do Estado de Minas Gerais (FAPEMIG \u0026ndash; APQ-01347-22).\u003c/p\u003e\u003ch2\u003eAvailability of data and material\u003c/h2\u003e \u003cp\u003eThe data will be made available for consultation, if required\u003c/p\u003e\u003ch2\u003eCode availability\u003c/h2\u003e \u003cp\u003eNot applicable\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAdams DG (2000) Symbiotic interactions. In: Whitton BA, Potts M (Eds.) The Ecology of Cyanobacteria, Their Diversity in Time and Space, Kluwer Academic Publishers, Dordrecht, the Netherlands, pp. 523\u0026ndash;61.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAguiar R, Fiore MF, Franco MW, Ventrella MC, Lorenzi AS, Vanetti CA, Alfenas AC (2008) A Novel epiphytic cyanobacterial species from the genus \u003cem\u003eBrasilonema\u003c/em\u003e causing damage to \u003cem\u003eEucalyptus\u003c/em\u003e leaves 1. Journal of Phycology, 44(5), 1322\u0026ndash;1334. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1529-8817.2008.00584.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1529-8817.2008.00584.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAhmed M, Stal LJ, Hasnain S (2010) Association of non-heterocystous cyanobacteria with crop plants. Plant and Soil, 336, 363\u0026ndash;375. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11104-010-0488-x\u003c/span\u003e\u003cspan address=\"10.1007/s11104-010-0488-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAlahuhta J, Lindholm M, Baastrup-Spohr L, Garc\u0026iacute;a-Gir\u0026oacute;n J, Toivanen M, Heino J, Murphy K (2021) Macroecology of macrophytes in the freshwater realm: Patterns, mechanisms and implications. Aquatic Botany, 168, 103325.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBiondi M (2021) Adapta\u0026ccedil;\u0026otilde;es de cianobact\u0026eacute;rias filamentosas e seus efeitos nas estrat\u0026eacute;gias reprodutivas de \u003cem\u003ePistia stratiotes\u003c/em\u003e. 69 f. Tese (Doutorado) - Curso de Bot\u0026acirc;nica Aplicada, Universidade Federal de Lavras.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBornette G, Puijalon S (2011) Response of aquatic plants to abiotic factors: a review. Aquatic sciences, 73, 1\u0026ndash;14. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00027-010-0162-7\u003c/span\u003e\u003cspan address=\"10.1007/s00027-010-0162-7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCameron HJ, Julian GR (1988) Utilization of hydroxyapatite by cyanobacteria as their sole source of phosphate and calcium, Plant and soil, 109, 123\u0026ndash;124. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/BF02197589\u003c/span\u003e\u003cspan address=\"10.1007/BF02197589\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCoelho FF, Basulto SR, F\u0026aacute;varo A, Demetrio GR (2021) Sex allocation in aquatic ferns: The role of body size. Plant Species Biol. 2021; 36: 425\u0026ndash;435. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/1442-1984.12324\u003c/span\u003e\u003cspan address=\"10.1111/1442-1984.12324\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCoelho FF, Deboni L, Santos Lopes F (2005a) Density-dependent reproductive and vegetative allocation in the aquatic plant \u003cem\u003ePistia stratiotes\u003c/em\u003e (Araceae), Revista de biolog\u0026iacute;a tropical, 53(3\u0026ndash;4), 369\u0026ndash;376. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.15517/rbt.v53i3-4.14599\u003c/span\u003e\u003cspan address=\"10.15517/rbt.v53i3-4.14599\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCoelho FF, Lopes FS, Sperber, CF (2005b) Persistence strategy of \u003cem\u003eSalvinia auriculata\u003c/em\u003e Aublet in temporary ponds of Southern Pantanal, Brazil, Aquatic Botany, 81(4), 343\u0026ndash;352. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.aquabot.2005.02.001\u003c/span\u003e\u003cspan address=\"10.1016/j.aquabot.2005.02.001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCrawley MJ (2012) The R book. John Wiley \u0026amp; Sons.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDar NA, Pandit AK, Ganai BA (2014) Factors Affecting the Distribution Patterns of Aquatic Macrophytes. Limnological Review, 14(2):75\u0026ndash;81. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2478/limre-2014-0008\u003c/span\u003e\u003cspan address=\"10.2478/limre-2014-0008\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDemetrio GR, Coelho FF (2023) What are the consequences of clonal integration for floral traits and reproductive investment of a broadly distributed aquatic plant? Flora, 303, 152292\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e.https://doi.org/10.1016/j.flora.2023.152292\u003c/span\u003e\u003cspan address=\".10.1016/j.flora.2023.152292\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDe Kroon H, Hutchings MJ (1995) Morphological plasticity in clonal plants: the foraging concept reconsidered, Journal of Ecology, 143\u0026ndash;152. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/2261158\u003c/span\u003e\u003cspan address=\"10.2307/2261158\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDe La Sota ER (1962) Contribuci\u0026oacute;n al conocimiento de Las Salviniaceae neotropicales, I \u003cem\u003eSalvinia oblongifolia\u003c/em\u003e Martius. Darwiniana, 465\u0026ndash;498.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDıez B, Ininbergs K (2014) Ecological importance of cyanobacteria. In: Sharma NK, Rai AK, Stal LJ (eds) Cyanobacteria: an economic perspective. Wiley, Chichester. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/9781118402238.ch3\u003c/span\u003e\u003cspan address=\"10.1002/9781118402238.ch3\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDong M, During HJ, Werger MJ (1997) Clonal plasticity in response to nutrient availability in the pseudoannual herb, \u003cem\u003eTrientalis europaea\u003c/em\u003e L, Plant Ecology, 131, 233\u0026ndash;239. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1023/A:1009783921753\u003c/span\u003e\u003cspan address=\"10.1023/A:1009783921753\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDroop MR (1968) Vitamin B12 and marine ecology. IV. The kinetics of uptake, growth and inhibition in \u003cem\u003eMonochrysis lutheri\u003c/em\u003e, Journal of the Marine Biological Association of the United Kingdom, 48(3), 689\u0026ndash;733. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1017/S0025315400019238\u003c/span\u003e\u003cspan address=\"10.1017/S0025315400019238\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDuan YH, Zhang YL, Ye LT, Fan XR, Xu, GH, Shen, QR (2007). Responses of rice cultivars with different nitrogen use efficiency to partial nitrate nutrition, Annals of Botany, 99(6), 1153\u0026ndash;1160. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/aob/mcm051\u003c/span\u003e\u003cspan address=\"10.1093/aob/mcm051\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eElbert W, Weber B, Burrows S, Steinkamp J, B\u0026uuml;del B, Andreae MO, P\u0026ouml;schl U (2012) Contribution of cryptogamic covers to the global cycles of carbon and nitrogen, Nature Geoscience, 5(7), 459\u0026ndash;462. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/ngeo1486\u003c/span\u003e\u003cspan address=\"10.1038/ngeo1486\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFranklin S, Alpert P, Salguero-G\u0026oacute;mez R, Janovsk\u0026yacute; Z, Herben T, Klimešov\u0026aacute; J, Douhovnikoff V (2021), Next-gen plant clonal ecology, Perspectives in plant ecology, evolution and systematics, 49, 125601. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ppees.2021.125601\u003c/span\u003e\u003cspan address=\"10.1016/j.ppees.2021.125601\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGadgil M, Bossert WH (1970) Life historical consequences of natural selection, The American Naturalist, 104(935), 1\u0026ndash;24. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1086/282637\u003c/span\u003e\u003cspan address=\"10.1086/282637\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGeng XM, He WM (2021) Success of native and invasive plant congeners depends on inorganic nitrogen compositions and levels, Journal of Plant Ecology, 14(2), 202\u0026ndash;212. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jpe/rtaa088\u003c/span\u003e\u003cspan address=\"10.1093/jpe/rtaa088\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGenu\u0026aacute;rio DB, Sant'Anna CL, Melo IS (2018) Elucidating the \u003cem\u003eCronbergia\u003c/em\u003e (Cyanobacteria) dilemma with the description of \u003cem\u003eCronbergia amazonensis\u003c/em\u003e sp. nov. isolated from Solim\u0026otilde;es river (Amazonia, Brazil), Algal research, 29, 233\u0026ndash;241. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.algal.2017.11.034\u003c/span\u003e\u003cspan address=\"10.1016/j.algal.2017.11.034\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGonz\u0026aacute;lez APR, Chrtek J, Dobrev, PI, Dumalasov\u0026aacute; V, Fehrer J, Mr\u0026aacute;z P, Latzel V (2016) Stress-induced memory alters growth of clonal offspring of white clover (\u003cem\u003eTrifolium repens\u003c/em\u003e), American Journal of Botany, 103(9), 1567\u0026ndash;1574. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3732/ajb.1500526\u003c/span\u003e\u003cspan address=\"10.3732/ajb.1500526\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGufu GD, Manea A, Leishman MR (2019) Growth, reproduction and functional trait responses of three freshwater plant species to elevated carbon dioxide, Aquatic Botany, 154, 18\u0026ndash;23. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.aquabot.2018.12.006\u003c/span\u003e\u003cspan address=\"10.1016/j.aquabot.2018.12.006\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHaroun SA, Hussein, MH (2003) The promotive effect of algal biofertilizers on growth, protein pattern and some metabolic activities of \u003cem\u003eLupinus termis\u003c/em\u003e plants grown in siliceous soil, Asian journal of Plant sciences. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3923/ajps.2003.944.951\u003c/span\u003e\u003cspan address=\"10.3923/ajps.2003.944.951\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHarper JL (1977) Population biology of plants, Population biology of plants.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHashtroudi MS, Ghassempour A, Riahi H, Shariatmadari Z, Khanjir M (2013) Endogenous auxins in plant growth-promoting Cyanobacteria\u0026mdash;\u003cem\u003eAnabaena vaginicola\u003c/em\u003e and \u003cem\u003eNostoc calcicola\u003c/em\u003e. Journal of applied phycology, 25, 379\u0026ndash;386. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10811-012-9872-7\u003c/span\u003e\u003cspan address=\"10.1007/s10811-012-9872-7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHempel M, Blume M, Blindow I, Gross EM (2008) Epiphytic bacterial community composition on two common submerged macrophytes in brackish water and freshwater, Bmc Microbiology, 8(1), 1\u0026ndash;10. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/1471-2180-8-58\u003c/span\u003e\u003cspan address=\"10.1186/1471-2180-8-58\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHoffman BM, Lukoyanov D, Yang ZY, Dean DR, Seefeldt LC (2014) Mechanism of nitrogen fixation by nitrogenase: the next stage. Chemical reviews, 114(8), 4041\u0026ndash;4062. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1021/cr400641x\u003c/span\u003e\u003cspan address=\"10.1021/cr400641x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHrouzek P, Lukešov\u0026aacute; A, Mareš J, Ventura S (2013) Description of the cyanobacterial genus \u003cem\u003eDesmonostoc\u003c/em\u003e gen. nov. including \u003cem\u003eD. muscorum\u003c/em\u003e comb. nov. as a distinct, phylogenetically coherent taxon related to the genus \u003cem\u003eNostoc\u003c/em\u003e, Fottea, 13(2), 201\u0026ndash;213. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.5507/fot.2013.016\u003c/span\u003e\u003cspan address=\"10.5507/fot.2013.016\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eJulien MH, Center TD, Tipping PW (2002) Floating fern (salvinia), Biological control of invasive plants in the Eastern United States, 17\u0026ndash;32.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKholssi R, Marks EA, Mi\u0026ntilde;\u0026oacute;n J, Mat\u0026eacute; AP, Sacrist\u0026aacute;n G, Montero O, Rad C (2021) A consortium of cyanobacteria and plant growth promoting rhizobacteria for wheat growth improvement in a hydroponic system, South African Journal of Botany, 142, 247\u0026ndash;258. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.sajb.2021.06.035\u003c/span\u003e\u003cspan address=\"10.1016/j.sajb.2021.06.035\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKluge M, Mollenhauer D, Wolf E, Sch\u0026uuml;\u0026szlig;ler A (2002) The \u003cem\u003eNostoc\u003c/em\u003e-\u003cem\u003eGeosiphon\u003c/em\u003e endocytobiosis, IN: Cyanobacteria in symbiosis, 19\u0026ndash;30.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKollah B, Patra AK, Mohanty SR (2016) Aquatic microphylla \u003cem\u003eAzolla\u003c/em\u003e: a perspective paradigm for sustainable agriculture, environment and global climate change, Environmental Science and Pollution Research, 23, 4358\u0026ndash;4369. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11356-015-5857-9\u003c/span\u003e\u003cspan address=\"10.1007/s11356-015-5857-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKollmen J, Strieth D (2022) The beneficial effects of cyanobacterial co-culture on plant growth, Life, 12(2), 223. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/life12020223\u003c/span\u003e\u003cspan address=\"10.3390/life12020223\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKom\u0026aacute;rek J (2016) A polyphasic approach for the taxonomy of cyanobacteria: principles and applications, European Journal of Phycology, 51(3), 346\u0026ndash;353. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/09670262.2016.1163738\u003c/span\u003e\u003cspan address=\"10.1080/09670262.2016.1163738\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKom\u0026aacute;rek J, Zapomelova E, Hindak F (2010) \u003cem\u003eCronbergia\u003c/em\u003e gen. nov., a new cyanobacterial genus (Cyanophyta) with a special strategy of heterocyte formation, Cryptogamie, Algologie, 31(3), 321\u0026ndash;341.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKom\u0026aacute;rek J, Johansen JR (2015) Filamentous cyanobacteria, In: Freshwater Algae of North America, Academic Press, 135\u0026ndash;235. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/B978-0-12-385876-4.00004-9\u003c/span\u003e\u003cspan address=\"10.1016/B978-0-12-385876-4.00004-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKuypers MM, Marchant HK, Kartal B (2018) The microbial nitrogen-cycling network, Nature Reviews Microbiology, 16(5), 263\u0026ndash;276. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/nrmicro.2018.9\u003c/span\u003e\u003cspan address=\"10.1038/nrmicro.2018.9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLacoul P, Freedman B (2006) Relationships between aquatic plants and environmental factors along a steep Himalayan altitudinal gradient. Aquat Bot 84:3\u0026ndash;16\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLefcheck JS (2016) \u0026ldquo;piecewiseSEM: Piecewise structural equation modeling in R for ecology, evolution, and systematics.\u0026rdquo; \u003cem\u003eMethods in Ecology and Evolution\u003c/em\u003e, 7(5), 573\u0026ndash;579. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/2041-210X.12512 \u0026lt;https://doi.org/10.1111/2041-210X.12512\u0026gt;\u003c/span\u003e\u003cspan address=\"10.1111/2041-210X.12512 %3C10.1111/2041-210X.12512%3E\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eManjunath M, Prasanna R, Sharma P, Nain L, Singh R (2011) Developing PGPR consortia using novel genera \u003cem\u003eProvidencia\u003c/em\u003e and \u003cem\u003eAlcaligenes\u003c/em\u003e along with cyanobacteria for wheat, Archives of Agronomy and Soil Science, 57(8), 873\u0026ndash;887. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/03650340.2010.499902\u003c/span\u003e\u003cspan address=\"10.1080/03650340.2010.499902\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMazhar S, Cohen JD, Hasnain S (2019) Novel Approach for the Determination of Nitrogen Fixation in Cyanobacteria, Journal of the Chemical Society of Pakistan, 41(1), 105\u0026ndash;105.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMedeiros JCC, Coelho, FF, Teixeira E (2016) Biomass allocation and nutrients balance related to the concentration of Nitrogen and Phosphorus in \u003cem\u003eSalvinia auriculata\u003c/em\u003e (Salviniaceae). Brazilian Journal of Biology, 76, 461\u0026ndash;468. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1590/1519-6984.21114\u003c/span\u003e\u003cspan address=\"10.1590/1519-6984.21114\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMedeiros JCC, Silva JCF, Resende TDSC, Teodoro GS, Pereira FJ, Coelho, FF (2018) Ramet versus sporocarp production in the aquatic fern \u003cem\u003eSalvinia auriculata\u003c/em\u003e (Salviniaceae): the role of shading, Australian Journal of Botany, 66(7), 583\u0026ndash;588. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1071/BT18062\u003c/span\u003e\u003cspan address=\"10.1071/BT18062\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMedeiros JCC, Silva JCF, Teodoro GS, Coelho FF (2017) Effects of shade on individual ramet growth and on clonal growth of the aquatic fern \u003cem\u003eSalvinia auriculata\u003c/em\u003e (Salviniaceae), American Fern Journal, 107(1), 21\u0026ndash;29. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1640/0002-8444-107.1.21\u003c/span\u003e\u003cspan address=\"10.1640/0002-8444-107.1.21\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMeeks JC (2005) Molecular mechanisms in the nitrogen-fixing \u003cem\u003eNostoc\u003c/em\u003e-bryophyte symbiosis, Molecular basis of symbiosis, 165\u0026ndash;196.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMeeks JC, Campbell EL, Summers ML, Wong FC (2002) Cellular differentiation in the cyanobacterium \u003cem\u003eNostoc punctiforme\u003c/em\u003e, Archives of microbiology, 178, 395\u0026ndash;403. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00203-002-0476-5\u003c/span\u003e\u003cspan address=\"10.1007/s00203-002-0476-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMiranda CV, Schwartsburd PB (2019) \u003cem\u003eSalvinia\u003c/em\u003e (Salviniaceae) in southern and southeastern Brazil\u0026mdash;including new taxa, new distribution records, and new morphological characters, Brazilian Journal of Botany, 42, 171\u0026ndash;188. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s40415-019-00522-5\u003c/span\u003e\u003cspan address=\"10.1007/s40415-019-00522-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMohamed ZA, Al Shehri AM (2010) Differential responses of epiphytic and planktonic toxic cyanobacteria to allelopathic substances of the submerged macrophyte \u003cem\u003eStratiotes aloides\u003c/em\u003e, International Review of Hydrobiology, 95(3), 224\u0026ndash;234. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/iroh.200911219\u003c/span\u003e\u003cspan address=\"10.1002/iroh.200911219\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eO\u0026rsquo;Hare MT, Baattrup-Pedersen A, Baumgarte I, Freeman A, Gunn ID, L\u0026aacute;z\u0026aacute;r AN, Bowes MJ (2018) Responses of aquatic plants to eutrophication in rivers: a revised conceptual model, Frontiers in plant science, 9, 451. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fpls.2018.00451\u003c/span\u003e\u003cspan address=\"10.3389/fpls.2018.00451\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOsman MEH, El-Sheekh MM, El-Naggar AH, Gheda SF (2010) Effect of two species of cyanobacteria as biofertilizers on some metabolic activities, growth, and yield of pea plant, Biology and fertility of soils, 46, 861\u0026ndash;875. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00374-010-0491-7\u003c/span\u003e\u003cspan address=\"10.1007/s00374-010-0491-7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eP\u0026eacute;rillon C, Hilt S (2019) Groundwater discharge gives periphyton a competitive advantage over macrophytes, Aquatic Botany, 154, 72\u0026ndash;80. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.aquabot.2019.01.004\u003c/span\u003e\u003cspan address=\"10.1016/j.aquabot.2019.01.004\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePrasanna R, Babu S, Rana A, Kabi SR, Chaudhary V, Gupta V, Pal RK (2013). Evaluating the establishment and agronomic proficiency of cyanobacterial consortia as organic options in wheat\u0026ndash;rice cropping sequence, Experimental agriculture, 49(3), 416\u0026ndash;434. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1017/S001447971200107X\u003c/span\u003e\u003cspan address=\"10.1017/S001447971200107X\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePrasanna R, Kanchan A, Ramakrishnan B, Ranjan K, Venkatachalam S, Hossain F, Nain L (2016a) Cyanobacteria-based bioinoculants influence growth and yields by modulating the microbial communities favourably in the rhizospheres of maize hybrids, European Journal of Soil Biology, 75, 15\u0026ndash;23. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ejsobi.2016.04.001\u003c/span\u003e\u003cspan address=\"10.1016/j.ejsobi.2016.04.001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePrasanna R, Ramakrishnan B, Ranjan K, Venkatachalam S, Kanchan A, Solanki P, Kranthi S (2016b) Microbial inoculants with multifaceted traits suppress \u003cem\u003eRhizoctonia\u003c/em\u003e populations and promote plant growth in cotton, Journal of Phytopathology, 164(11\u0026ndash;12), 1030\u0026ndash;1042. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/jph.12524\u003c/span\u003e\u003cspan address=\"10.1111/jph.12524\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePrasanna R, Ramakrishnan B, Simranjit K, Ranjan K, Kanchan A, Hossain F, Nain L (2017) Cyanobacterial and rhizobial inoculation modulates the plant physiological attributes and nodule microbial communities of chickpea, Archives of Microbiology, 199, 1311\u0026ndash;1323. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00203-017-1405-y\u003c/span\u003e\u003cspan address=\"10.1007/s00203-017-1405-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePimenta LL, Lima GP, Biondi M, Vaz MGMV, Coelho FF (2022) Epiphytic cyanobacterial strains in the roots of \u003cem\u003eSalvinia auriculata\u003c/em\u003e and the effect of light and nutrients on the production of heterocyst, akinete and hormogonia, Aquatic Ecology, 56(3), 543\u0026ndash;553. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10452-021-09922-w\u003c/span\u003e\u003cspan address=\"10.1007/s10452-021-09922-w\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePott VJ, Pott A (2000) Plantas aqu\u0026aacute;ticas do Pantanal, Bras\u0026iacute;lia, DF: Embrapa Comunica\u0026ccedil;\u0026atilde;o para Transfer\u0026ecirc;ncia de Tecnologia; Corumb\u0026aacute;: Embrapa-CPAP.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRai AN, Bergman B, Rasmussen U (2002) Cyanobacteria in symbiosis, Kluwer Academic Pub.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRai AN, Singh AK, Syiem MB (2019) Plant growth-promoting abilities in cyanobacteria, In: Cyanobacteria, Academic Press, 459\u0026ndash;476\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eR Core Team (2023) \u003cem\u003eR: A Language and Environment for Statistical Computing\u003c/em\u003e. R Foundation for Statistical Computing, Vienna, Austria. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e\u0026lt;https://www.R-project.org/\u0026gt;\u003c/span\u003e\u003cspan address=\"http://%3Chttps://www.R-project.org/%3E\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eŘeh\u0026aacute;kov\u0026aacute; K, Johansen JR, Casamatta DA, Xuesong L, Vincent J (2007) Morphological and molecular characterization of selected desert soil cyanobacteria: three species new to science including \u003cem\u003eMojavia pulchra\u003c/em\u003e gen. et sp. nov.,Phycologia, 46(5), 481\u0026ndash;502. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2216/06-92.1\u003c/span\u003e\u003cspan address=\"10.2216/06-92.1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRolon AS, Maltchik L (2006) Environmental factors as predictors of aquatic macrophyte richness and composition in wetlands of southern Brazil. Hydrobiologia, 556, 221\u0026ndash;231.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRomo S (2007) Epiphyton, phytoplankton and macrophyte ecology in a shallow lake under in situ experimental conditions, Fundamental and applied limnology, 197\u0026ndash;209. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1127/1863-9135/2007/0170-0197\u003c/span\u003e\u003cspan address=\"10.1127/1863-9135/2007/0170-0197\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRoom PM (1983) Falling apart'as a lifestyle: the rhizome architecture and population growth of \u003cem\u003eSalvinia molesta\u003c/em\u003e, The Journal of Ecology, 349\u0026ndash;365. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/2259719\u003c/span\u003e\u003cspan address=\"10.2307/2259719\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSadvakasova AK, Bauenova MO, Kossalbayev BD, Zayadan BK, Huang Z, Wang J, Allakhverdiev SI (2023) Synthetic algocyanobacterial consortium as an alternative to chemical fertilizers, Environmental Research, 116418. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.envres.2023.116418\u003c/span\u003e\u003cspan address=\"10.1016/j.envres.2023.116418\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSant\u0026rsquo;anna CL, Branco LHZ, Azevedo MTP, Werner VR, Malone CFS (2020) Cyanobacteria, In: Bicudo CEM, Menezes M, G\u0026ecirc;neros de Algas de \u0026Aacute;guas Continentais no Brasil (4) p. 33\u0026ndash;41.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSantamar\u0026iacute;a L (2002) Why are most aquatic plants widely distributed? Dispersal, clonal growth and small-scale heterogeneity in a stressful environment. Acta oecologica, 23(3), 137\u0026ndash;154.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSchmitt J, Flemming HC (1999) Water binding in biofilms, Water science and technology, 39(7), 77\u0026ndash;82. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/S0273-1223(99)00153-5\u003c/span\u003e\u003cspan address=\"10.1016/S0273-1223(99)00153-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSchooling SR, Beveridge TJ (2006) Membrane vesicles: an overlooked component of the matrices of biofilms, Journal of bacteriology, 188(16), 5945\u0026ndash;5957. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1128/jb.00257-06\u003c/span\u003e\u003cspan address=\"10.1128/jb.00257-06\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSculthorpe CD (1967) The biology of aquatic vascular plants.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSeastedt T (2009) Traits of plant invaders, Nature, 459(7248), 783\u0026ndash;784. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/459783a\u003c/span\u003e\u003cspan address=\"10.1038/459783a\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSergeeva E, Liaimer A, Bergman B (2002) Evidence for production of the phytohormone indole-3-acetic acid by cyanobacteria, Planta, 215, 229\u0026ndash;238. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00425-002-0749-x\u003c/span\u003e\u003cspan address=\"10.1007/s00425-002-0749-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eShariatmadari Z, Riahi H, Seyed Hashtroudi M, Ghassempour A, Aghashariatmadary Z (2013) Plant growth promoting cyanobacteria and their distribution in terrestrial habitats of Iran, Soil Science and Plant Nutrition, 59(4), 535\u0026ndash;547. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/00380768.2013.782253\u003c/span\u003e\u003cspan address=\"10.1080/00380768.2013.782253\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSingha A (2009) Diversity of Cyanobacteria in Organic Farming Field Under Rice-wheat Cropping System (Doctoral dissertation, IARI, Division of Microbiology, New Delhi).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSmith SD (2014) The roles of nitrogen and phosphorus in regulating the dominance of floating and submerged aquatic plants in a field mesocosm experiment, Aquatic Botany, 112, 1\u0026ndash;9. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.aquabot.2013.07.001\u003c/span\u003e\u003cspan address=\"10.1016/j.aquabot.2013.07.001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSrivastava JK, Chandra H, Kalra SJ, Mishra P, Khan H, Yadav P (2017) Plant\u0026ndash;microbe interaction in aquatic system and their role in the management of water quality: a review, Applied Water Science, 7, 1079\u0026ndash;1090. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s13201-016-0415-2\u003c/span\u003e\u003cspan address=\"10.1007/s13201-016-0415-2\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eStewart WDP, Rowell P, Rai AN (1983) Cyanobacteria-eukaryotic plant symbioses, In: Annales de l'Institut Pasteur/Microbiologie, Elsevier Masson 134 (1) 205\u0026ndash;228. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/S0769-2609(83)80106-9\u003c/span\u003e\u003cspan address=\"10.1016/S0769-2609(83)80106-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVerma S, Adak A, Prasanna R, Dhar S, Choudhary H, Nain L, Shivay YS (2016) Microbial priming elicits improved plant growth promotion and nutrient uptake in pea, Israel Journal of Plant Sciences, 63(3), 191\u0026ndash;207.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang R, Peng B, Huang K (2015) The research progress of CO\u0026lt;background-color:#CCCCFF;subdirection:rtl;\u0026gt;2\u0026lt;/background-color:#CCCCFF;subdirection:rtl;\u0026gt; sequestration by algal bio-fertilizer in China, Journal of CO\u0026lt;background-color:#CCFF99;subdirection:rtl;\u0026gt;2\u0026lt;/background-color:#CCFF99;subdirection:rtl;\u0026gt; Utilization, 11, 67\u0026ndash;70. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.jcou.2015.01.007\u003c/span\u003e\u003cspan address=\"10.1016/j.jcou.2015.01.007\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eXie D, Yu D, You WH, Wang LG (2013) Algae mediate submerged macrophyte response to nutrient and dissolved inorganic carbon loading: a mesocosm study on different species, Chemosphere, 93(7), 1301\u0026ndash;1308. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.chemosphere.2013.07.008\u003c/span\u003e\u003cspan address=\"10.1016/j.chemosphere.2013.07.008\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYandigeri MS, Meena KK, Srinivasan R, Pabbi S (2011) Effect of mineral phosphate solubilization on biological nitrogen fixation by diazotrophic cyanobacteria, Indian journal of microbiology, 51, 48\u0026ndash;53. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s12088-011-0081-x\u003c/span\u003e\u003cspan address=\"10.1007/s12088-011-0081-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang LM, Jin Y, Yao SM, Lei NF, Chen JS, Zhang Q, Yu, FH (2020) Growth and morphological responses of duckweed to clonal fragmentation, nutrient availability, and population density, Frontiers in Plant Science, 11, 618.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZheng Z, Bai W, Zhang WH (2019) Clonality-dependent dynamic change of plant community in temperate grasslands under nitrogen enrichment, Oecologia, 189, 255\u0026ndash;266. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00442-018-4317-x\u003c/span\u003e\u003cspan address=\"10.1007/s00442-018-4317-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhou X, Wang G, Yang F (2011) Characteristics of growth, nutrient uptake, purification effect of \u003cem\u003eIpomoea aquatica\u003c/em\u003e, \u003cem\u003eLolium multiflorum\u003c/em\u003e, and \u003cem\u003eSorghum sudanense\u003c/em\u003e grown under different nitrogen levels, Desalination, 273(2\u0026ndash;3), 366\u0026ndash;374. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.desal.2011.01.057\u003c/span\u003e\u003cspan address=\"10.1016/j.desal.2011.01.057\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZuo S, Liu L, Zhou C, Wang X, Huang D (2023) Effects of nitrogen limitation on \u003cem\u003eLeymus chinensis\u003c/em\u003e clonal growth form and bud bank, Flora, (298) 152186. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.flora.2022.152186\u003c/span\u003e\u003cspan address=\"10.1016/j.flora.2022.152186\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Aquatic Plant, Biofertiliser, Clonal Growth, Heterocystous Cyanobacteria, Reproductive Strategies","lastPublishedDoi":"10.21203/rs.3.rs-5822876/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5822876/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eClonal plants, like \u003cem\u003eSalvinia auriculata\u003c/em\u003e, are widespread and perform important ecosystem functions, influencing the structure and composition of the ecosystems in which they occur. Some cyanobacteria perform biological nitrogen fixation (BNF) and can affect plant growth, as nitrogen (N) is a limiting nutrient. Therefore, we carried out a greenhouse experiment with the inoculation of the cyanobacterial strain \u003cem\u003eDesmonostoc\u003c/em\u003e (UFLA 12) to investigate whether heterocystous cyanobacteria favour individual growth and drives reproductive strategies (sexual reproduction and/or clonal growth) of \u003cem\u003eS. auriculata\u003c/em\u003e. \u003cem\u003eSalvinia auriculata\u003c/em\u003e ramets were grown in plastic pots under the following treatments: (D) \u003cem\u003eDesmonostoc \u003c/em\u003e(UFLA 12) inoculum and (Co) control, in which cyanobacteria were absent. The \u003cem\u003eDesmonostoc \u003c/em\u003epresence positively influenced the clonal growth of \u003cem\u003eS. auriculata\u003c/em\u003e, and increased shoots number, plants fresh biomass, and shoot size. We conclude that the inoculation of \u003cem\u003eDesmonostoc \u003c/em\u003e(UFLA 12) contributes to a more vigorous spread of \u003cem\u003eS. auriculata\u003c/em\u003e since it enhanced clonal growth. In this sense, the role of \u003cem\u003eDesmonostoc \u003c/em\u003e(UFLA 12) as a potential biofertilizer may serve as a tool to assist in understanding the excessive growth of \u003cem\u003e\u0026nbsp;S. auriculata\u003c/em\u003e in aquatic environments.\u003c/p\u003e","manuscriptTitle":"Cyanobacterial inoculation promotes enhanced clonal growth performance of the aquaticplant Salvinia auriculata.","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-01-17 15:40:47","doi":"10.21203/rs.3.rs-5822876/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"b20450c8-8f70-4ede-afbf-583bdf14d58d","owner":[],"postedDate":"January 17th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-02-11T12:23:41+00:00","versionOfRecord":[],"versionCreatedAt":"2025-01-17 15:40:47","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-5822876","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5822876","identity":"rs-5822876","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}