Effects of high pH on a natural marine planktonic community

Effects of high pH on a natural marine planktonic community Effects of high pH on a natural marine planktonic community

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22 Mar Ecol Prog Ser 260: 19–31, 2003 Fig. 2. Fluctuations in nutrient concentrations measured during the 2 wk experimental period. (A) NH 4 + , (B) SiO4 – , (C) NO3 – + NO 2 – , (D) PO4 2– . (y) ong>pHong> 8.0; (j)ong>pHong> 8.5; (S) ong>pHong> 9.0; (m) ong>pHong> 9.5. Symbols represent means ong>ofong> triplicates ± SE Fig. 3. Phytoplankton concentrations in the 4 incubations during the 2 wk experimental period. (A) Chlorophyll a, (B) phytoplankton biomass (µg C l –1 ), (C) diatom biomass, (D) phototrophic dinong>ofong>lagellate biomass. (y) ong>pHong> 8.0; (j) ong>pHong> 8.5; (S) ong>pHong> 9.0; (m) ong>pHong> 9.5. Symbols represent means ong>ofong> triplicates ± SE

Pedersen & Hansen: ong>Effectsong> ong>ofong> ong>highong> ong>pHong> on a planktonic community grew throughout the duration ong>ofong> the experiment, whereas the dinong>ofong>lagellates only grew for the first 3 d. At ong>highong>er ong>pHong>, the biomass ong>ofong> both diatoms and dinong>ofong>lagellates either remained constant or declined during the first week ong>ofong> the incubation, and only growth ong>ofong> diatoms was observed during the second week ong>ofong> the incubations. The succession among the species within the studied phytoplankton groups varied according to the ong>pHong> level (Figs. 4 & 5). The 2 lowest ong>pHong> incubations, ong>pHong> 8 and 8.5, experienced almost no succession among species. Here, all the identified species were present throughout the experimental period. In the ong>pHong> 9.5 incubation, a pronounced succession ong>ofong> species occurred during the incubation period. At the end ong>ofong> the experiment, Cylindrotheca closterium was the only species among the diatoms that thrived. Its growth rate at ong>pHong> 9.5 was similar to the growth rates obtained in the lower ong>pHong> incubations (Fig. 4, Table 1). Among the dinong>ofong>lagellates, Prorocentrum micans, P. minimum and Heterocapsa triquetra all survived at ong>pHong> 9.5, whereas the initial dominant species, Ceratium furca, C. fusus and C. tripos, died out (Fig. 5). In the ong>pHong> 9 incubation, the succession ong>ofong> species was less pronounced, and only a few species died out. It was interesting to note however that some species apparently grew faster in the ong>pHong> 9 incubation compared to in the incubations at lower ong>pHong> (Figs. 4 & 5). Protozooplankton At the start ong>ofong> the experiment, the protozooplankton biomass was 20 µg C l –1 in all incubations (Fig. 6A). In the ong>pHong> 8, 8.5 and 9 incubations, a general increase in biomass was found over time. An 8-fold increase in biomass was observed at the termination ong>ofong> the experiment in the ong>pHong> 8 and 9 incubations, whereas only a 3-fold increase in biomass was found in the ong>pHong> 8.5 incubation. In the ong>pHong> 9.5 incubation, the biomass decreased about 5-fold during the first 3 d and stayed at that level until the termination ong>ofong> the experiment. The increase in biomass in the ong>pHong> 8, 8.5 and 9 incubations was mainly caused by ciliates, because the heterotrophic dinong>ofong>lagellates were found to be relatively constant throughout the experiment. In the ong>pHong> 9.5 incubation, both ciliates and heterotrophic dinong>ofong>lagellates declined in biomass throughout the duration ong>ofong> the experiment (Fig. 6B,C). The succession among the species within the studied protozooplankton groups varied according to the ong>pHong> level (Figs. 7 & 8). The 2 lowest ong>pHong> incubations, ong>pHong> 8 and 8.5, experienced almost no succession among species. Here, all the identified species were present throughout the experimental period. In the ong>pHong> 9.5 Fig. 4. Cell concentration ong>ofong> some selected diatoms in the 4 incubations during the 2 wk experimental period. (A) Cylindrotheca closterium, (B) Cerataulina pelagica, (C) Leptocylindrus minimus. (y) ong>pHong> 8.0; (j) ong>pHong> 8.5; (S) ong>pHong> 9.0; (m) ong>pHong> 9.5. Symbols represent means ong>ofong> triplicates ± SE incubation, some species died out, whereas the remaining species were alive at the termination ong>ofong> the experiment. However, unlike in the case ong>ofong> the phytoplankton, none ong>ofong> the protozooplankton species took over. In the ong>pHong> 9 incubation, the succession ong>ofong> species was less pronounced, and only a few species died out (Figs. 7 & 8). It was interesting to note however that some ciliate species apparently grew faster in the ong>pHong> 9 incubation compared to in the incubations at lower ong>pHong>. 23

Pedersen & Hansen: <str<strong>on</strong>g>Effects</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>high</str<strong>on</strong>g> <str<strong>on</strong>g>pH</str<strong>on</strong>g> <strong>on</strong> a plankt<strong>on</strong>ic <strong>community</strong><br />

grew throughout the durati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> the experiment,<br />

whereas the din<str<strong>on</strong>g>of</str<strong>on</strong>g>lagellates <strong>on</strong>ly grew for the first 3 d.<br />

At <str<strong>on</strong>g>high</str<strong>on</strong>g>er <str<strong>on</strong>g>pH</str<strong>on</strong>g>, the biomass <str<strong>on</strong>g>of</str<strong>on</strong>g> both diatoms and din<str<strong>on</strong>g>of</str<strong>on</strong>g>lagellates<br />

either remained c<strong>on</strong>stant or declined during<br />

the first week <str<strong>on</strong>g>of</str<strong>on</strong>g> the incubati<strong>on</strong>, and <strong>on</strong>ly growth <str<strong>on</strong>g>of</str<strong>on</strong>g><br />

diatoms was observed during the sec<strong>on</strong>d week <str<strong>on</strong>g>of</str<strong>on</strong>g> the<br />

incubati<strong>on</strong>s.<br />

The successi<strong>on</strong> am<strong>on</strong>g the species within the studied<br />

phytoplankt<strong>on</strong> groups varied according to the <str<strong>on</strong>g>pH</str<strong>on</strong>g> level<br />

(Figs. 4 & 5). The 2 lowest <str<strong>on</strong>g>pH</str<strong>on</strong>g> incubati<strong>on</strong>s, <str<strong>on</strong>g>pH</str<strong>on</strong>g> 8 and<br />

8.5, experienced almost no successi<strong>on</strong> am<strong>on</strong>g species.<br />

Here, all the identified species were present throughout<br />

the experimental period.<br />

In the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9.5 incubati<strong>on</strong>, a pr<strong>on</strong>ounced successi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g><br />

species occurred during the incubati<strong>on</strong> period. At the<br />

end <str<strong>on</strong>g>of</str<strong>on</strong>g> the experiment, Cylindrotheca closterium was<br />

the <strong>on</strong>ly species am<strong>on</strong>g the diatoms that thrived. Its<br />

growth rate at <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9.5 was similar to the growth rates<br />

obtained in the lower <str<strong>on</strong>g>pH</str<strong>on</strong>g> incubati<strong>on</strong>s (Fig. 4, Table 1).<br />

Am<strong>on</strong>g the din<str<strong>on</strong>g>of</str<strong>on</strong>g>lagellates, Prorocentrum micans, P.<br />

minimum and Heterocapsa triquetra all survived at<br />

<str<strong>on</strong>g>pH</str<strong>on</strong>g> 9.5, whereas the initial dominant species, Ceratium<br />

furca, C. fusus and C. tripos, died out (Fig. 5).<br />

In the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9 incubati<strong>on</strong>, the successi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> species was<br />

less pr<strong>on</strong>ounced, and <strong>on</strong>ly a few species died out. It<br />

was interesting to note however that some species<br />

apparently grew faster in the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9 incubati<strong>on</strong> compared<br />

to in the incubati<strong>on</strong>s at lower <str<strong>on</strong>g>pH</str<strong>on</strong>g> (Figs. 4 & 5).<br />

Protozooplankt<strong>on</strong><br />

At the start <str<strong>on</strong>g>of</str<strong>on</strong>g> the experiment, the protozooplankt<strong>on</strong><br />

biomass was 20 µg C l –1 in all incubati<strong>on</strong>s (Fig. 6A). In<br />

the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 8, 8.5 and 9 incubati<strong>on</strong>s, a general increase in<br />

biomass was found over time. An 8-fold increase in<br />

biomass was observed at the terminati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> the experiment<br />

in the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 8 and 9 incubati<strong>on</strong>s, whereas <strong>on</strong>ly a<br />

3-fold increase in biomass was found in the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 8.5<br />

incubati<strong>on</strong>. In the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9.5 incubati<strong>on</strong>, the biomass<br />

decreased about 5-fold during the first 3 d and stayed<br />

at that level until the terminati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> the experiment.<br />

The increase in biomass in the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 8, 8.5 and 9 incubati<strong>on</strong>s<br />

was mainly caused by ciliates, because the<br />

heterotrophic din<str<strong>on</strong>g>of</str<strong>on</strong>g>lagellates were found to be relatively<br />

c<strong>on</strong>stant throughout the experiment. In the<br />

<str<strong>on</strong>g>pH</str<strong>on</strong>g> 9.5 incubati<strong>on</strong>, both ciliates and heterotrophic<br />

din<str<strong>on</strong>g>of</str<strong>on</strong>g>lagellates declined in biomass throughout the<br />

durati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> the experiment (Fig. 6B,C).<br />

The successi<strong>on</strong> am<strong>on</strong>g the species within the studied<br />

protozooplankt<strong>on</strong> groups varied according to the <str<strong>on</strong>g>pH</str<strong>on</strong>g><br />

level (Figs. 7 & 8). The 2 lowest <str<strong>on</strong>g>pH</str<strong>on</strong>g> incubati<strong>on</strong>s, <str<strong>on</strong>g>pH</str<strong>on</strong>g> 8<br />

and 8.5, experienced almost no successi<strong>on</strong> am<strong>on</strong>g<br />

species. Here, all the identified species were present<br />

throughout the experimental period. In the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9.5<br />

Fig. 4. Cell c<strong>on</strong>centrati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> some selected diatoms in the 4 incubati<strong>on</strong>s<br />

during the 2 wk experimental period. (A) Cylindrotheca<br />

closterium, (B) Cerataulina pelagica, (C) Leptocylindrus<br />

minimus. (y) <str<strong>on</strong>g>pH</str<strong>on</strong>g> 8.0; (j) <str<strong>on</strong>g>pH</str<strong>on</strong>g> 8.5; (S) <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9.0; (m) <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9.5.<br />

Symbols represent means <str<strong>on</strong>g>of</str<strong>on</strong>g> triplicates ± SE<br />

incubati<strong>on</strong>, some species died out, whereas the remaining<br />

species were alive at the terminati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> the<br />

experiment. However, unlike in the case <str<strong>on</strong>g>of</str<strong>on</strong>g> the phytoplankt<strong>on</strong>,<br />

n<strong>on</strong>e <str<strong>on</strong>g>of</str<strong>on</strong>g> the protozooplankt<strong>on</strong> species took<br />

over.<br />

In the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9 incubati<strong>on</strong>, the successi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> species was<br />

less pr<strong>on</strong>ounced, and <strong>on</strong>ly a few species died out<br />

(Figs. 7 & 8). It was interesting to note however that<br />

some ciliate species apparently grew faster in the <str<strong>on</strong>g>pH</str<strong>on</strong>g> 9<br />

incubati<strong>on</strong> compared to in the incubati<strong>on</strong>s at lower <str<strong>on</strong>g>pH</str<strong>on</strong>g>.<br />

23

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