Allelopathic effects of the aquatic macrophyte Ceratophyllum demersum L. on phytoplankton species: contrasting effects between cyanobacteria and chlorophytes

Abstract Aim To assess the allelopathic effects of the submerged macrophyte Ceratophyllum demersum on four strains of phytoplankton species: two cyanobacteria (Microcystis aeruginosa - microcystin producing and M. panniformis - non-microcystin producing), and two chlorophytes (Ankistrodesmus falcatu...

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Veröffentlicht in:Acta limnológica brasiliensia 2019, Vol.31, p.1-10
Hauptverfasser: Amorim, Cihelio Alves, Moura-Falcão, Rafael Henrique de, Valença, Celina Rebeca, Souza, Vitor Ricardo de, Moura, Ariadne do Nascimento
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Valença, Celina Rebeca
Souza, Vitor Ricardo de
Moura, Ariadne do Nascimento
description Abstract Aim To assess the allelopathic effects of the submerged macrophyte Ceratophyllum demersum on four strains of phytoplankton species: two cyanobacteria (Microcystis aeruginosa - microcystin producing and M. panniformis - non-microcystin producing), and two chlorophytes (Ankistrodesmus falcatus and Raphidocelis subcapitata). Methods A coexistence experiment between C. demersum and the four strains was carried out for six days, with eight treatments and three replicates. The strains were cultivated in ASM1 culture medium, under controlled laboratory conditions. Two treatments were assigned for each strain, one with 6 g.L-1 of the macrophyte, and the control without the plant. Biomasses and growth rates of the strains were evaluated every two days, which were compared through the T-test and two-way ANOVA, respectively. Results The results varied among the strains, with toxic M. aeruginosa being intensely inhibited by C. demersum, with a decrease of 99.5% in its biomass (p
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Methods A coexistence experiment between C. demersum and the four strains was carried out for six days, with eight treatments and three replicates. The strains were cultivated in ASM1 culture medium, under controlled laboratory conditions. Two treatments were assigned for each strain, one with 6 g.L-1 of the macrophyte, and the control without the plant. Biomasses and growth rates of the strains were evaluated every two days, which were compared through the T-test and two-way ANOVA, respectively. Results The results varied among the strains, with toxic M. aeruginosa being intensely inhibited by C. demersum, with a decrease of 99.5% in its biomass (p<0.001), while non-toxic M. panniformis was less affected by the allelochemicals, with a reduction of 86.2% (p<0.001). Ankistrodesmus falcatus delayed its growth when in coexistence with the macrophyte, decreasing its biomass in 50.4% (p<0.01), while R. subcapitata was not altered (p>0.05). In coexistence with C. demersum, M. aeruginosa exhibited the lowest growth rates (-0.65 d-1), followed by M. panniformis (-0.15 d-1), A. falcatus (0.19 d-1), and R. subcapitata (0.34 d-1), with significant differences between all strains (p<0.001). Microcystis aeruginosa presented higher inhibition rates than M. panniformis (p<0.001), as well as, A. falcatus was more inhibited than R. subcapitata (p<0.05). Conclusions The presence of microcystins could influence the allelopathic responses of C. demersum, that may release more allelochemicals in coexistence with toxic strains of M. aeruginosa. Accordingly, C. demersum can be used in biomanipulation strategies to control toxic and non-toxic cyanobacterial blooms, without damaging other phytoplankton species, like chlorophytes. Resumo Objetivo Avaliar os efeitos alelopáticos da macrófita submersa Ceratophyllum demersum sobre quatro cepas de espécies fitoplanctônicas: duas cianobactérias (Microcystis aeruginosa - produtora de microcistinas e M. panniformis - não produtora) e duas clorófitas (Ankistrodesmus falcatus e Raphidocelis subcapitata). Métodos Foi realizado um experimento de coexistência entre C. demersum e as quatro cepas, durante seis dias, com oito tratamentos e três réplicas. As cepas foram cultivadas em meio ASM1, sob condições laboratoriais controladas. Foram designados dois tratamentos para cada cepa, um com 6 g.L-1 da macrófita, e um controle sem a planta. Biomassas e taxas de crescimento das cepas foram avaliadas a cada dois dias e comparadas pelo teste-T e ANOVA two-way, respectivamente. Resultados Os resultados variaram entre as cepas, sendo a cepa tóxica de M. aeruginosa intensamente inibida por C. demersum, com uma redução de 99,5% na sua biomassa (p<0,001), enquanto a cepa não tóxica de M. panniformis foi menos afetada pelos aleloquímicos, com uma redução de 86,2% (p<0,001). Ankistrodesmus falcatus retardou seu crescimento em coexistência com a macrófita, reduzindo sua biomassa em 50,4% (p<0,01), enquanto que R. subcapitata não foi afetada (p>0,05). Em coexistência com C. demersum, M. aeruginosa apresentou as menores taxas de crescimento (-0,65 d-1), seguida de M. panniformis (-0,15 d-1), A. falcatus (0,19 d-1) e R. subcapitata (0,34 d-1), com diferenças significativas entre todas as cepas (p<0,001). Microcystis aeruginosa apresentou maiores taxas de inibição que Microcystis panniformis (p<0,001), bem como, A. falcatus foi mais inibida que R. subcapitata (p<0,05). Conclusões A presença de microcistinas pode afetar as respostas alelopáticas de C. demersum, que pode liberar mais aleloquímicos em coexistência com cepas tóxicas de M. aeruginosa. Portanto, C. demersum pode ser utilizada em estratégias de biomanipulação para controle de florações de cianobactérias tóxicas e não tóxicas, sem causar danos às demais espécies fitoplanctônicas, como as clorófitas.]]></description><identifier>ISSN: 2179-975X</identifier><identifier>ISSN: 0102-6712</identifier><identifier>EISSN: 2179-975X</identifier><identifier>DOI: 10.1590/s2179-975x1419</identifier><language>eng</language><publisher>Botucatu: Associação Brasileira de Limnologia</publisher><subject>alelopatia ; Algae ; Allelochemicals ; Allelopathy ; Ankistrodesmus falcatus ; Aquatic plants ; Biomanipulation ; biomanipulação ; Biomass ; Blooms ; Ceratophyllum demersum ; Coexistence ; controle de cianobactérias ; Culture media ; Cyanobacteria ; Ecology ; experimentos de coexistência ; Freshwater plants ; Growth rate ; Laboratories ; Lakes ; LIMNOLOGY ; macrófitas submersas ; Microcystins ; Microcystis ; Microcystis aeruginosa ; Phytoplankton ; Plankton ; Raphidocelis subcapitata ; Species ; Toxicology ; Toxins ; Variance analysis</subject><ispartof>Acta limnológica brasiliensia, 2019, Vol.31, p.1-10</ispartof><rights>2019. This work is published under https://creativecommons.org/licenses/by/4.0 (the “License”). Notwithstanding the ProQuest Terms and Conditions, you may use this content in accordance with the terms of the License.</rights><rights>This work is licensed under a Creative Commons Attribution 4.0 International License.</rights><lds50>peer_reviewed</lds50><oa>free_for_read</oa><woscitedreferencessubscribed>false</woscitedreferencessubscribed><citedby>FETCH-LOGICAL-c3939-266dc2df6a25b6ad31305f1f29e430335f03c0d33a35a35e034e65f30255c9a53</citedby><cites>FETCH-LOGICAL-c3939-266dc2df6a25b6ad31305f1f29e430335f03c0d33a35a35e034e65f30255c9a53</cites><orcidid>0000-0002-7171-7450 ; 0000-0001-5093-2840</orcidid></display><links><openurl>$$Topenurl_article</openurl><openurlfulltext>$$Topenurlfull_article</openurlfulltext><thumbnail>$$Tsyndetics_thumb_exl</thumbnail><link.rule.ids>230,314,776,780,860,881,4009,27902,27903,27904</link.rule.ids></links><search><creatorcontrib>Amorim, Cihelio Alves</creatorcontrib><creatorcontrib>Moura-Falcão, Rafael Henrique de</creatorcontrib><creatorcontrib>Valença, Celina Rebeca</creatorcontrib><creatorcontrib>Souza, Vitor Ricardo de</creatorcontrib><creatorcontrib>Moura, Ariadne do Nascimento</creatorcontrib><title>Allelopathic effects of the aquatic macrophyte Ceratophyllum demersum L. on phytoplankton species: contrasting effects between cyanobacteria and chlorophytes</title><title>Acta limnológica brasiliensia</title><addtitle>Acta Limnol. Bras</addtitle><description><![CDATA[Abstract Aim To assess the allelopathic effects of the submerged macrophyte Ceratophyllum demersum on four strains of phytoplankton species: two cyanobacteria (Microcystis aeruginosa - microcystin producing and M. panniformis - non-microcystin producing), and two chlorophytes (Ankistrodesmus falcatus and Raphidocelis subcapitata). Methods A coexistence experiment between C. demersum and the four strains was carried out for six days, with eight treatments and three replicates. The strains were cultivated in ASM1 culture medium, under controlled laboratory conditions. Two treatments were assigned for each strain, one with 6 g.L-1 of the macrophyte, and the control without the plant. Biomasses and growth rates of the strains were evaluated every two days, which were compared through the T-test and two-way ANOVA, respectively. Results The results varied among the strains, with toxic M. aeruginosa being intensely inhibited by C. demersum, with a decrease of 99.5% in its biomass (p<0.001), while non-toxic M. panniformis was less affected by the allelochemicals, with a reduction of 86.2% (p<0.001). Ankistrodesmus falcatus delayed its growth when in coexistence with the macrophyte, decreasing its biomass in 50.4% (p<0.01), while R. subcapitata was not altered (p>0.05). In coexistence with C. demersum, M. aeruginosa exhibited the lowest growth rates (-0.65 d-1), followed by M. panniformis (-0.15 d-1), A. falcatus (0.19 d-1), and R. subcapitata (0.34 d-1), with significant differences between all strains (p<0.001). Microcystis aeruginosa presented higher inhibition rates than M. panniformis (p<0.001), as well as, A. falcatus was more inhibited than R. subcapitata (p<0.05). Conclusions The presence of microcystins could influence the allelopathic responses of C. demersum, that may release more allelochemicals in coexistence with toxic strains of M. aeruginosa. Accordingly, C. demersum can be used in biomanipulation strategies to control toxic and non-toxic cyanobacterial blooms, without damaging other phytoplankton species, like chlorophytes. Resumo Objetivo Avaliar os efeitos alelopáticos da macrófita submersa Ceratophyllum demersum sobre quatro cepas de espécies fitoplanctônicas: duas cianobactérias (Microcystis aeruginosa - produtora de microcistinas e M. panniformis - não produtora) e duas clorófitas (Ankistrodesmus falcatus e Raphidocelis subcapitata). Métodos Foi realizado um experimento de coexistência entre C. demersum e as quatro cepas, durante seis dias, com oito tratamentos e três réplicas. As cepas foram cultivadas em meio ASM1, sob condições laboratoriais controladas. Foram designados dois tratamentos para cada cepa, um com 6 g.L-1 da macrófita, e um controle sem a planta. Biomassas e taxas de crescimento das cepas foram avaliadas a cada dois dias e comparadas pelo teste-T e ANOVA two-way, respectivamente. Resultados Os resultados variaram entre as cepas, sendo a cepa tóxica de M. aeruginosa intensamente inibida por C. demersum, com uma redução de 99,5% na sua biomassa (p<0,001), enquanto a cepa não tóxica de M. panniformis foi menos afetada pelos aleloquímicos, com uma redução de 86,2% (p<0,001). Ankistrodesmus falcatus retardou seu crescimento em coexistência com a macrófita, reduzindo sua biomassa em 50,4% (p<0,01), enquanto que R. subcapitata não foi afetada (p>0,05). Em coexistência com C. demersum, M. aeruginosa apresentou as menores taxas de crescimento (-0,65 d-1), seguida de M. panniformis (-0,15 d-1), A. falcatus (0,19 d-1) e R. subcapitata (0,34 d-1), com diferenças significativas entre todas as cepas (p<0,001). Microcystis aeruginosa apresentou maiores taxas de inibição que Microcystis panniformis (p<0,001), bem como, A. falcatus foi mais inibida que R. subcapitata (p<0,05). Conclusões A presença de microcistinas pode afetar as respostas alelopáticas de C. demersum, que pode liberar mais aleloquímicos em coexistência com cepas tóxicas de M. aeruginosa. Portanto, C. demersum pode ser utilizada em estratégias de biomanipulação para controle de florações de cianobactérias tóxicas e não tóxicas, sem causar danos às demais espécies fitoplanctônicas, como as clorófitas.]]></description><subject>alelopatia</subject><subject>Algae</subject><subject>Allelochemicals</subject><subject>Allelopathy</subject><subject>Ankistrodesmus falcatus</subject><subject>Aquatic plants</subject><subject>Biomanipulation</subject><subject>biomanipulação</subject><subject>Biomass</subject><subject>Blooms</subject><subject>Ceratophyllum demersum</subject><subject>Coexistence</subject><subject>controle de cianobactérias</subject><subject>Culture media</subject><subject>Cyanobacteria</subject><subject>Ecology</subject><subject>experimentos de coexistência</subject><subject>Freshwater plants</subject><subject>Growth rate</subject><subject>Laboratories</subject><subject>Lakes</subject><subject>LIMNOLOGY</subject><subject>macrófitas submersas</subject><subject>Microcystins</subject><subject>Microcystis</subject><subject>Microcystis aeruginosa</subject><subject>Phytoplankton</subject><subject>Plankton</subject><subject>Raphidocelis subcapitata</subject><subject>Species</subject><subject>Toxicology</subject><subject>Toxins</subject><subject>Variance analysis</subject><issn>2179-975X</issn><issn>0102-6712</issn><issn>2179-975X</issn><fulltext>true</fulltext><rsrctype>article</rsrctype><creationdate>2019</creationdate><recordtype>article</recordtype><sourceid>ABUWG</sourceid><sourceid>AFKRA</sourceid><sourceid>AZQEC</sourceid><sourceid>BENPR</sourceid><sourceid>CCPQU</sourceid><sourceid>DWQXO</sourceid><sourceid>DOA</sourceid><recordid>eNpNUk1vGyEQXVWt1CjJtWeknu0Cs-ya3CIrTSNZ6iGp1Bsas0O8Ll42gNX4x_S_lo3zhUDMjOa9GXhTVV8Enwul-bckRatnulWPohb6Q3Xy4v_--M7-XJ2ntOVlLRa1rPVJ9e_Se_JhxLzpLSPnyObEgmN5Qwwf9phLeIc2hnFzyMSWFDFPtvf7HetoRzEVYzVnYWBTShg9Dn9y8dJItqd0wWwYcsSU--H-tcKa8l-igdkDDmGNNlPskeHQMbvx4blaOqs-OfSJzp_v0-rX96u75Y_Z6uf1zfJyNbOgQc9k03RWdq5BqdYNdiCAKyec1FQDB1COg-UdAIIqmzjU1CgHXCplNSo4rW6OvF3ArRljv8N4MAF78xQI8d5gLD_hyaBuOEpEEqqt0XXaFR5a61ZI7copXPMjVyqv98Fswz4OpXlzO8lgJhkkF7poIDgHsSiAr0fAGMPDnlJ-g0jZQtOqhVRvtEWKlCK51zYFN9MImPTC_zQC8B8JHaUD</recordid><startdate>2019</startdate><enddate>2019</enddate><creator>Amorim, Cihelio Alves</creator><creator>Moura-Falcão, Rafael Henrique de</creator><creator>Valença, Celina Rebeca</creator><creator>Souza, Vitor Ricardo de</creator><creator>Moura, Ariadne do Nascimento</creator><general>Associação Brasileira de Limnologia</general><scope>AAYXX</scope><scope>CITATION</scope><scope>7QH</scope><scope>7QL</scope><scope>7SN</scope><scope>7SS</scope><scope>7UA</scope><scope>ABUWG</scope><scope>AEUYN</scope><scope>AFKRA</scope><scope>AZQEC</scope><scope>BENPR</scope><scope>BHPHI</scope><scope>BKSAR</scope><scope>C1K</scope><scope>CCPQU</scope><scope>CLZPN</scope><scope>DWQXO</scope><scope>F1W</scope><scope>H95</scope><scope>H96</scope><scope>HCIFZ</scope><scope>L.G</scope><scope>M7N</scope><scope>PCBAR</scope><scope>PIMPY</scope><scope>PQEST</scope><scope>PQQKQ</scope><scope>PQUKI</scope><scope>GPN</scope><scope>DOA</scope><orcidid>https://orcid.org/0000-0002-7171-7450</orcidid><orcidid>https://orcid.org/0000-0001-5093-2840</orcidid></search><sort><creationdate>2019</creationdate><title>Allelopathic effects of the aquatic macrophyte Ceratophyllum demersum L. on phytoplankton species: contrasting effects between cyanobacteria and chlorophytes</title><author>Amorim, Cihelio Alves ; Moura-Falcão, Rafael Henrique de ; Valença, Celina Rebeca ; Souza, Vitor Ricardo de ; Moura, Ariadne do Nascimento</author></sort><facets><frbrtype>5</frbrtype><frbrgroupid>cdi_FETCH-LOGICAL-c3939-266dc2df6a25b6ad31305f1f29e430335f03c0d33a35a35e034e65f30255c9a53</frbrgroupid><rsrctype>articles</rsrctype><prefilter>articles</prefilter><language>eng</language><creationdate>2019</creationdate><topic>alelopatia</topic><topic>Algae</topic><topic>Allelochemicals</topic><topic>Allelopathy</topic><topic>Ankistrodesmus falcatus</topic><topic>Aquatic plants</topic><topic>Biomanipulation</topic><topic>biomanipulação</topic><topic>Biomass</topic><topic>Blooms</topic><topic>Ceratophyllum demersum</topic><topic>Coexistence</topic><topic>controle de cianobactérias</topic><topic>Culture media</topic><topic>Cyanobacteria</topic><topic>Ecology</topic><topic>experimentos de coexistência</topic><topic>Freshwater plants</topic><topic>Growth rate</topic><topic>Laboratories</topic><topic>Lakes</topic><topic>LIMNOLOGY</topic><topic>macrófitas submersas</topic><topic>Microcystins</topic><topic>Microcystis</topic><topic>Microcystis aeruginosa</topic><topic>Phytoplankton</topic><topic>Plankton</topic><topic>Raphidocelis subcapitata</topic><topic>Species</topic><topic>Toxicology</topic><topic>Toxins</topic><topic>Variance analysis</topic><toplevel>peer_reviewed</toplevel><toplevel>online_resources</toplevel><creatorcontrib>Amorim, Cihelio Alves</creatorcontrib><creatorcontrib>Moura-Falcão, Rafael Henrique de</creatorcontrib><creatorcontrib>Valença, Celina Rebeca</creatorcontrib><creatorcontrib>Souza, Vitor Ricardo de</creatorcontrib><creatorcontrib>Moura, Ariadne do Nascimento</creatorcontrib><collection>CrossRef</collection><collection>Aqualine</collection><collection>Bacteriology Abstracts (Microbiology B)</collection><collection>Ecology Abstracts</collection><collection>Entomology Abstracts (Full archive)</collection><collection>Water Resources Abstracts</collection><collection>ProQuest Central (Alumni Edition)</collection><collection>ProQuest One Sustainability</collection><collection>ProQuest Central UK/Ireland</collection><collection>ProQuest Central Essentials</collection><collection>ProQuest Central</collection><collection>Natural Science Collection (ProQuest)</collection><collection>Earth, Atmospheric &amp; 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Bras</addtitle><date>2019</date><risdate>2019</risdate><volume>31</volume><spage>1</spage><epage>10</epage><pages>1-10</pages><issn>2179-975X</issn><issn>0102-6712</issn><eissn>2179-975X</eissn><abstract><![CDATA[Abstract Aim To assess the allelopathic effects of the submerged macrophyte Ceratophyllum demersum on four strains of phytoplankton species: two cyanobacteria (Microcystis aeruginosa - microcystin producing and M. panniformis - non-microcystin producing), and two chlorophytes (Ankistrodesmus falcatus and Raphidocelis subcapitata). Methods A coexistence experiment between C. demersum and the four strains was carried out for six days, with eight treatments and three replicates. The strains were cultivated in ASM1 culture medium, under controlled laboratory conditions. Two treatments were assigned for each strain, one with 6 g.L-1 of the macrophyte, and the control without the plant. Biomasses and growth rates of the strains were evaluated every two days, which were compared through the T-test and two-way ANOVA, respectively. Results The results varied among the strains, with toxic M. aeruginosa being intensely inhibited by C. demersum, with a decrease of 99.5% in its biomass (p<0.001), while non-toxic M. panniformis was less affected by the allelochemicals, with a reduction of 86.2% (p<0.001). Ankistrodesmus falcatus delayed its growth when in coexistence with the macrophyte, decreasing its biomass in 50.4% (p<0.01), while R. subcapitata was not altered (p>0.05). In coexistence with C. demersum, M. aeruginosa exhibited the lowest growth rates (-0.65 d-1), followed by M. panniformis (-0.15 d-1), A. falcatus (0.19 d-1), and R. subcapitata (0.34 d-1), with significant differences between all strains (p<0.001). Microcystis aeruginosa presented higher inhibition rates than M. panniformis (p<0.001), as well as, A. falcatus was more inhibited than R. subcapitata (p<0.05). Conclusions The presence of microcystins could influence the allelopathic responses of C. demersum, that may release more allelochemicals in coexistence with toxic strains of M. aeruginosa. Accordingly, C. demersum can be used in biomanipulation strategies to control toxic and non-toxic cyanobacterial blooms, without damaging other phytoplankton species, like chlorophytes. Resumo Objetivo Avaliar os efeitos alelopáticos da macrófita submersa Ceratophyllum demersum sobre quatro cepas de espécies fitoplanctônicas: duas cianobactérias (Microcystis aeruginosa - produtora de microcistinas e M. panniformis - não produtora) e duas clorófitas (Ankistrodesmus falcatus e Raphidocelis subcapitata). Métodos Foi realizado um experimento de coexistência entre C. demersum e as quatro cepas, durante seis dias, com oito tratamentos e três réplicas. As cepas foram cultivadas em meio ASM1, sob condições laboratoriais controladas. Foram designados dois tratamentos para cada cepa, um com 6 g.L-1 da macrófita, e um controle sem a planta. Biomassas e taxas de crescimento das cepas foram avaliadas a cada dois dias e comparadas pelo teste-T e ANOVA two-way, respectivamente. Resultados Os resultados variaram entre as cepas, sendo a cepa tóxica de M. aeruginosa intensamente inibida por C. demersum, com uma redução de 99,5% na sua biomassa (p<0,001), enquanto a cepa não tóxica de M. panniformis foi menos afetada pelos aleloquímicos, com uma redução de 86,2% (p<0,001). Ankistrodesmus falcatus retardou seu crescimento em coexistência com a macrófita, reduzindo sua biomassa em 50,4% (p<0,01), enquanto que R. subcapitata não foi afetada (p>0,05). Em coexistência com C. demersum, M. aeruginosa apresentou as menores taxas de crescimento (-0,65 d-1), seguida de M. panniformis (-0,15 d-1), A. falcatus (0,19 d-1) e R. subcapitata (0,34 d-1), com diferenças significativas entre todas as cepas (p<0,001). Microcystis aeruginosa apresentou maiores taxas de inibição que Microcystis panniformis (p<0,001), bem como, A. falcatus foi mais inibida que R. subcapitata (p<0,05). Conclusões A presença de microcistinas pode afetar as respostas alelopáticas de C. demersum, que pode liberar mais aleloquímicos em coexistência com cepas tóxicas de M. aeruginosa. Portanto, C. demersum pode ser utilizada em estratégias de biomanipulação para controle de florações de cianobactérias tóxicas e não tóxicas, sem causar danos às demais espécies fitoplanctônicas, como as clorófitas.]]></abstract><cop>Botucatu</cop><pub>Associação Brasileira de Limnologia</pub><doi>10.1590/s2179-975x1419</doi><tpages>10</tpages><orcidid>https://orcid.org/0000-0002-7171-7450</orcidid><orcidid>https://orcid.org/0000-0001-5093-2840</orcidid><oa>free_for_read</oa></addata></record>
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subjects alelopatia
Algae
Allelochemicals
Allelopathy
Ankistrodesmus falcatus
Aquatic plants
Biomanipulation
biomanipulação
Biomass
Blooms
Ceratophyllum demersum
Coexistence
controle de cianobactérias
Culture media
Cyanobacteria
Ecology
experimentos de coexistência
Freshwater plants
Growth rate
Laboratories
Lakes
LIMNOLOGY
macrófitas submersas
Microcystins
Microcystis
Microcystis aeruginosa
Phytoplankton
Plankton
Raphidocelis subcapitata
Species
Toxicology
Toxins
Variance analysis
title Allelopathic effects of the aquatic macrophyte Ceratophyllum demersum L. on phytoplankton species: contrasting effects between cyanobacteria and chlorophytes
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