Diet selection by copepods in the presence of cyanobacteria

Boeckella, the dominant calanoid in many Southern Hemispherelakes, can survive, grow and reproduce to varying extents onmonocultures of cyanobacteria. In this study, we determinedthe effects of algal and cyanobacterial foods of different nutritionalvalue and concentration on food preferences of adult femaleBoeckella trianiculata and Boeckella hamata. Four species ofcyanobacteria (Anabaena flos-aquae, Nostoc sp. 2, OscillatoriatenuisandMicrocystis aeruginosa) were offered alone and mixedwith equal biomasses of Cryptomonas sp., Choricystis or a cyanobacterium.Food preferences were calculated as ratios of the rates at whichthe copepods removed each food at high and low food concentrations.In high-concentration mixtures with cyanobacteria, Cryptomonaswas consistently preferred by both Boeckella spp. In low-concentrationmixtures, both Boeckella spp. preferred Anabaena and Nostoc,which they removed at high rates(81–142 ml mg^–1h^–1), although Cryptomonas was selected in preferenceto Oscillatoria and Microcystis. When fed mixtures of filamentouscyanobacteria, both species of Boeckella showed invariant discriminationagainst Nostoc, andshifts in preference between Anabaena andOscillatoria that were related to food concentration. Microcystis,the least favouredfood, appeared to have a toxic effect on B.triarticulata. ¹Present address: Nursing and Midwifery Department, Otago Polytechnic,Forth Street, Dunedin, New Zealand     

Effects of food quality on individual growth and development in the freshwater copepod Boeckella triarticulata

To quantify the demographic effects of food quality, and specificallyof the ‘poorquality’ cyanobecterium Anabaena flos-aquae,we reared individual Boeckella triarticulata (Copepoda, Calanoida)on two diets (monospecific Cryptomom sp. versus mixed Cryptomonas-Anabanmadiets) and quantified individual growth and developmental trajectoriesby examining exuviae produced at each molt, from hatching tomaturity. Size at molting was less variable among individuals,within and between diets, than age. Food quality had significanteffects on male sizes at molting and on stage-specific dailygrowth rates of both sexes; these effects were strongest duringlate naupliar and all copepodite stages Tke med Cryptomonas-Anabaenadiet significantly slowed development, particularly of copepoditestages. As a consequence of these effects, individuals raisedon the mixed diet were smaller and older at maturity. Withina given diet, individual differences explained much, if notmost, of the variation exhibited in growth and development.By following growth and development of a large number of individualsthroughout their Life cycles, we show that individual femalesproduce variable offspring, indicative of a bet-hedging life-historystrategy, and that B.triarticulata (like other calanoids) cangrow, develop and survive on diets that include ‘poorquality’ cyanobacteria. ¹Present address: Department of Biological Sciences, Universityof Rhode Island, Kingston, RI 02881, USA     

The influence of food resources on the development, survival and reproduction of the two cyclopoid copepods: Cyclops vicinus and Mesocyclops leuckarti

Laboratory experiments were conducted to determine whether thetwo cyclopoid copepods. Cyclops vicious and Mesocyclops leuckaru.exploit the same food resources. The food requirements of juvenilesof the two cyclopoid copepods were investigated. Moreover, theimportance of algae for the predaceous adults was studied. Naupliiof both M leuckaru and C.vicinus successfully developed intocopepodites when fed the motile algae Chlamydomonas reinhardtii.Chlamydomonas sphaeroides and Cryptomonas sp. Threshold foodconcentrations for naupliar development varied between offeredalgae and between the two cyclopoid species. The food thresholdfor successful naupliar development, when reared on C.reinhardui,was lower for M.leuckarti (0.3 mg C 1^–1) than for C.vicinus(0.5 mg C l^–1) whereas a similar food threshold was foundusing Cryptornonas sp (0.3 mg C ^–1) and C.sphaeroides(<0.2 mg C 1^–1), Naupliar development time was inverselyrelated to food concentration. Food required for copepoditedevelopment differed for the two cyclopoid species. Cyclopsvicinus was able to develop to the adult stage on a pure dietof any one of the three algal species. whereas M.leuckarti requireda prey supply of the rotifer Brachionus rubens. Food composition.i.e. algal species, algal concentration and rotifer abundance,influenced copepodite survivorship of both cyclopoids and wasalways higher in the presence of B.rubens. Under similar foodconditions, mortality was higher for M.leuckarti than for Cvicinus. Mesocyclops leuckaru females were very dependent onanimal food. The predation rate of M.leuckaru was not lowerin the presence of algae. Egg production of M.leuckarti waslow on a pure algal diet and significantly higher when B rubenswas present. The results were used to discuss the life cyclestrategy and the possibility of exploitative competition ofthe two cyclopoid copepods.     

Effects of rotifers and ciliates on the growth and survival of Daphnia

Daphnia can suppress ciliates and rotifers through predationand interference competition, but it is not known whether thisproduces any direct benefit to Daphnia. We conducted survivorshipand cohort lifetable experiments to determine whether Daphniacan utilize ciliates and rotifers as food. Three species ofoligotrich ciliates (Halteria grandinella, Strobilidium gyransand Strobilidiumvelox) and one rotifer (Keratella cochlearis)were used. Lifetable experiments were conducted with a basallevel of algae (Cryptomonas sp.), plus either ciliates or rotifers,while survivorship experiments had only the rotifers or ciliates.Densities of 30 H.grandinella ml^–1, 50 S.gyrans ml^–1and 15 S.velox ml^–1 enhanced Daphnia pulex's populationgrowth rate 35–50% over controls with only algae. TenS.gyrans ml^–1 did not produce a significant change inDaphnia's growth rate. Densities of 100 and 300 K.cochlearis^–1 increased Daphnia population growth rates by II and10%, respectively. Both 10 and 50 S.gyrans ml^–1 enhancedDaphnia's survivorship compared to starved controls, but neither100 nor 300 K.cochlearis l^–1 enhanced its survivorship.The amount of enhancement of Daphnia growth rates by rotifersand ciliates is roughly proportional to the death rates imposedby Daphnia. The death rate imposed by Daphnia on rotifers isa function of both algal density and Daphnia size. Per unitbiomass, neither ciliates nor Keratella appear to be as nutritiousfor Daphnia as is Cryptomonas.     

Genetic diversity in cyanobacterial symbionts of thalloid bryophytes

Two species of thalloid liverworts, Blasia pusilla and Caviculariadensa, form stable symbioses with nitrogen-fixing cyanobacteria.Both bryophytes promote the persistence of their cyanobacterialassociations by producing specialized gemmae, which facilitatethe simultaneous dispersal of the host and its nitrogen-fixingsymbionts. Here the genetic diversity of cyanobacterial symbiontsof Blasia and Cavicularia is examined. The results indicatethat the primary symbionts of both bryophytes are closely relatedand belong to a specific group of symbiotic Nostoc strains.Related strains have previously been reported from hornwortsand cycads, and from many terricolous cyanolichens. The evolutionaryorigins of all these symbioses may trace back to pre-Permiantimes. While the laboratory strain Nostoc punctiforme PCC 73102has been widely used in experimental studies of bryophyte–Nostocassociations, sequence-identical cyanobionts have not yet beenidentified from thalloid liverworts in the field. Key words: Blasia pusilla, bryophyte, Cavicularia densa, Nostoc, tRNA^Leu(UAA) intron, specificity, symbiosis Received 27 May 2007; Revised 15 November 2007 Accepted 21 December 2007     

Predation by ciliates on a metalimnetic Cryptomonas population: feeding rates, impact and effects of vertical migration

Deep algal maxima are frequently overlayed by dense populationsof ciliates, rotifers and crustaceans. This has been interpretedas evidence of heavy predation on the algae, although the impactof this predation has never been determined experimentally.We determined the vertical and seasonal distribution of thealga Cryptomonas phaseolus and its most relevant predators,the ciliates Coleps sp. and Prorodon sp., forming metalimneticmaxima in Lake Cisó. On several dates, in situ feedingrates of ciliates were determined by three independent methods:(i) epifluorescence counts of ingested algal cells togetherwith estimates of the food turnover time of the ciliates; (ii)in situ incubations with radioactively labeled algae: (iii)HPLC determination of alloxanthine content in the predator sizefraction. Feeding rates varied between 0.07 and 0.64 Cryptomonasciliate^–1 h^–1. We then calculated integrated predationon the algae. using the functional response of the ciliatesand the vertical distribution of each population. We found thateven though the ciliates were always food saturated, their predationimpact on Cryptomonas was not very large: as an average, 5–25%of the biomass of Cryptomonas was removed daily by the ciliates.Finally, we studied the effects of the diel vertical movementsof these populations on predation impact. By migrating intothe sulfide-rich hypolimnion during the night, Cryptomonas couldreduce its predation losses by 38%. Thus, the algae were protectedfrom predation during several hours of each diel cycle and maintaineda very large biomass throughout stratification, although thisresulted in a very slow growth. Slow growth, coupled to largebiomass, seems to be a general feature of metalimnetic accumulationsof organisms.     

Relative impacts of copepods, cladocerans and nutrients on the microbial food web of a mesotrophic lake

Calanoid copepods, rather than cladocerans, frequently dominatethe zooplankton of lakes in New Zealand. The potential consequencesof this domination for the microbial community of mesotrophicLake Mahinerangi, New Zealand, were determined by field experimentsin which Boeckella and Daphnia were added to in situ enclosuresin the presence and absence of added nutrients. Boeckella hamataat ambient densities (2 and 81^–1) rapidly and severelysuppressed ciliate population growth over 4 days, even whenmicrobial growth was enhanced by added nutrients, but effectsof copepods on other components of the microbial community (bacteria,photosynthetic picoplankton, heterotrophic nano-flagellates,algae) were slight. In contrast, Daphnia carinata at the samedensities (but 3-fold higher biomasses per litre) had a relativelyweak effect on ciliates, suppressing ciliate abundance onlyafter 4 days at 8 Daphnia 1^–1 (330 µg 1^–1);this daphniid density also depressed abundances of large bacterialrods, some photosynthetic picoplankton and the dominant alga,Cyclotella. These results highlight the relative importanceof specific trophic linkages in a microbial food web; they alsosuggest that the dominance of Boeckella in many southern hemispherelakes may account for relatively low ciliate abundances in theselakes.     

Effect of Sodium on Phosphate Uptake in Unicellular and Filamentous Cyanobacteria

The effect of Na⁺ on phosphate uptake was studied in four strainsof cyanobacteria: Synechococcus PCC 7942, Gloeothece PCC 6501,Phormidium sp. and Chlorogloeopsis PCC 6912. Phosphate uptakewas stimulated by Na⁺ in all cases. Li⁺ and K⁺ acted as partialanalogues for Na⁺. Half-saturation [K_1/2(Na⁺)] of phosphateuptake was reached with Na⁺ concentrations ranging from 317µM in Chlorogloeopsis to 659 µM in Phormidium. Theconcentration of phosphate required to reach half-saturationof phosphate uptake [K_1/2(P_i)]was not changed by the presenceof Na⁺. (Received April 11, 1994; Accepted July 5, 1994)     

An Examination of some Symbiotic Systems for Fixation of Nitrogen

By the use of ¹⁵N the occurrence of fixation of elemental nitrogenhas been demonstrated in the lichens Collema granosum and Leptogiumlichenoides (both of which contain Nostoc as a symbiont), andalso in the liverwort Blasia pusilla (which has Nostoc-containingcavities in the thallus). The circumstances indicate that thefixation in these instances should be attributed to the Nostoc. No evidence of fixation was obtained in similar isotopic testson the mycorrhizal roots of intact plants of Calluna vulgarisand of Pinus sylvestris, although these, more especially inthe case of Calluna, have been held by some previous authorsto be nitrogen-fixing.     

Phytoplankton population dynamics in perennially ice-covered Lake Fryxell, Antarctica

Phytoplankton were collected over five austral summers (1987–88through 1991–92) to examine seasonal and annual fluctuationsin species composition and biovolume in Lake Fryxell, a perenniallyice-covered lake located in the Dry Valleys of South VictoriaLand, Antarctica. Lake Fryxell has perennial gradients of salinity,dissolved oxygen and nutrients. We found that algal speciesdiversity was low (56 taxa were collected), confirming the resultsof previous short-term studies. The phytoplankton consistedprimarily of cryptophyte and chlorophyte flagellates, and filamentouscyanobacteria. The presence of filamentous cyanobacteria, whichhave not been reported as abundant in this lake by previousworkers, may represent a significant ecological change. Eachaustral summer, one dominant species contributed >70% ofthe total biovolume; Chroomonas lacustris was dominant in 1987–88,while Cryptomonas sp. dominated the phytoplankton in the remaining4 years. No species succession occurred during the austral summer.Some common taxa were vertically stratified (Oscillatoria limnetica,Phormidium angustissimum, Pyramimonas sp., Oscillatoria sp.),while others showed no distinct vertical stratification (Chlamydomonassubcaudata, Cryptomonas sp.). The stratification of the phytoplanktonreflects the gradients of nutrients and light, and the stabilityof the water column.     

Phytoplankton seasonal dynamics in a Mediterranean coastal lagoon: emphasis on the picoeukaryote community

The dynamics of the phytoplankton community were investigatedin a marine coastal lagoon (Thau, NW Mediterranean) from February1999 to January 2000. Dilution experiments, chlorophyll a (Chla) size-fractionation and primary production measurements wereconducted monthly. Maximum growth and microzooplankton grazingrates were estimated from Chl a biomass fractions to separatepico- from nano- and microphytoplankton and by flow cytometryto distinguish between picoeukaryotes and picocyanobacteria.In spring, the phytoplankton community was dominated by Chaetocerossp. and Skeletonema costatum, which represented most of biomass(B) and primary production (P). Nano- and microphytoplanktongrowth was controlled by nutrient availability and exceededlosses due to microzooplankton grazing (g). Picoeukaryote andcyanobacteria growth was positively correlated with water temperatureand/or irradiance, reaching maximum values in the summer (2.38and 1.44 day^–1 for picoeukaryotes and cyanobacteria, respectively).Picophytoplankton accounted for 57% of the biomass-specificprimary productivity (P/B). Picophytoplankton was strongly controlledby protist grazers (g = 0.09–1.66 day^–1 for picoeukaryotes,g = 0.25–1.17 day^–1 for cyanobacteria), and microzooplanktonconsumption removed 71% of the daily picoplanktonic growth.Picoeukaryotes, which numerically dominate the picoplanktoncommunity, are an important source of organic carbon for theprotistan community and contribute to the carbon flow to highertrophic levels.     

Relationships between fecal pellet production and feeding in the calanoid copepod Boeckella

The number, size (mm³ pellet ^–1) and total volume of fecalpellets produced (mm³ h^–1) by Boeckella titicacae increasedwith concentration of Chlorella and natural seston in threedifferent lakewaters from Bahia de Puno and Lago Grande of LakeTiticaca and nearby Laguna Arapa. The three lakewaters differedin the size and number of fecal pellets produced at seston concentrations>0.5 mg dry wt l^–1. However, the total volume of fecalpellets egested varied little. Large-sized food (>10 µg)resulted in larger pellets but smaller total fecal production.Food concentration had parallel effects on the production offeces and on feeding rates. Both fecal production and feedingincreased linearly with food concentration with saturation between5 and 7.5 x 10⁵ Chlorella ml^–1 (3.7–5.6 mg dry wtl^–1). Varied responses in fecal production when fed differentsize fractions of lake seston suggest Boeckella has a complexand flexible feeding behavior. Feeding preferences of B.titicacaeand B.occidentalis (Ivlev index) for two seston size fractionsare generally reflected by fecal production.     

The fauna of athalassic saline waters in Australia and the Altiplano of South America: comparisons and historical perspectives

Similarities and differences between the fauna of inland saline waters in Australia and on the Altiplano are explored and explanations sought.Elements common to both continents include the calanoid copepod genus Boeckella (B. triarticulata in Australia, B. poopoensis and B. meteoris in South America) and the cladoceran genus Daphniopsis. Salinity data for Altiplano lakes are given for six species of Boeckella and for Daphniopsis.Ostracods have adapted to the open water of saline lakes in Australia but not in South America, a difference that may reflect past differences in the degree of predation by birds. In South America, diatoms are grazed by the flamingos Phoenicoparrus andinus and P. jamesi, while in Australia the main diatom grazer is probably the aquatic oniscoid isopod Haloniscus searlei. However, at least four species of flamingos were present in Australia during the late Cenozoic and one or more of these may well have grazed diatoms. The extinction of diatom-grazing or carnivorous flamingos, or both, in Australia may have been factors in the unique colonization of inland saline waters by H. searlei.     

In vitro Regeneration from Excised Leaf Discs of Three Brassica Species

Excised leaf discs of three Brassica species, B. oleracea, B.napus, and B. campestris were induced to produce adventitiousbuds and subsequently entire plants by culture on media withspecific combinations of 6-benzylaminopurine (BAP) and -naphthylaceticacid (NAA). Each species required a particular hormone concentrationfor optimum growth and differentiation: B. oleracea, BAP 10mg^–1 and NAA 1 mg 1^–1; B. napus, BAP 10 mg 1^–1and NAA 10 mg 1^–1; B. campestris, BAP 1 mg 1^–1 andNAA 10 mg 1^–1. In a more detailed study on one of these species, namely B.oleracea, the relative influence of other media components suchas amino acids and other organic additives was examined. Itwas also found that the source and size of the explant greatlyaffected the growth response, as did the size of the culturevessel. The regenerated plants dislayed a range of ploidy as well asphenotypic abnormalities. Findings are discussed in relation to results from other tissueculture systems.     

Experimental studies into the feeding biology of rotifers in brackish water

Mass developments of rotifers of the genus Brachionus, and especiallyof B.quadridentatus, occur regularly in the largely hypertrophicchain of shallow waters (‘boddens’) south of theDarss-Zingst peninsula (Southern Baltic). Interest in the autecologyof the species is, therefore, considerable. Various food sourceswere used in laboratory experiments to ascertain the food requirementsof B.quadridentatus, determine its filtration and ingestionrates, and assess its food particle size-selection ability.Growth experiments showed that the chlorophyceans Nannochlorissp. and Chlorella vulgaris possess considerable nutritionalvalue for the species, whereas abundances declined when Microcystisfirma, detritus from Enteromorpha sp. and only bacteria (Pseudomonas),respectively, were provided as food sources. Filtration ratesvaried between 0.02 and 1.73 µl ind.^–1 h^–1,and ingestion rates between 121 and 5560 cells ind.^–1h^–1, depending on the filtration rate and algal concentration.Investigations into food particle size selection using fluorescentlatex particles revealed that particle size influences foodparticle intake. When particles of different sizes were mixed,the animals showed a preference for the larger particles andingested the smaller ones with a diameter of 1–2 µmless efficiently. The brackish water species Brachionus plicatiliswas studied besides B.quadridentatus in all experiments. Theformer species proved to be superior both in its range of utilizableparticle sizes and its growth rate. The experiments with laboratorycultures were backed up by studies performed with various rotiferspecies taken from natural populations.     

Impact of Daphnia pulex on a metalimnetic microbial community

A well-structured metalimnetic community, composed mainly ofphototrophic purple bacteria, Cryptomonas, and a few speciesof ciliates and rotifers, was regularly found between 1984 and1991 in Lake Cis. These populations appeared during spring,reached high concentrations and remained in the metalimnionuntil winter mixing. We had previously postulated that thesemetalimnetic populations could persist in such high numbersbecause they were free of efficient predation, since the oxygen-sulfideinterface provided a refuge from predation. In September 1992,an abundant Daphnia pulex population developed in the lake whichresulted in drastic changes in the formerly stable community.The presence of D.pulex induced an extreme clear-water phasein the epilimnion and the metalimnetic populations were severelyreduced. Feeding experiments with Cryptomonas and heterotrophicand phototrophic bacteria as food for D.pulex resulted in clearancerates around 0.7 ml individual^–1 h^–1. The impactwas very high on the microaerophilic populations, whereas theanaerobic populations were least affected, since sulfide limitedthe vertical distribution of ^D.pulex. We conclude from thisevidence that the metalimnetic populations in previous yeanwere mostly food limited (bottom-up control) and that theirhigh abundances were in great part due to a lack of efficientpredation.     

Feeding interactions of the copepods Eurytemora affinis and Acartia bifilosa with the cyanobacteria Nodularia sp.

We measured ingestion and clearance rates of two Baltic Seacalanoid copepods, Eurytemora affinis and Acartia bifilosa,on toxic and non-toxic cyanobacteria Nodularia sp. using theisotope technique. Eurytemora affinis fed actively on the non-toxicstrain and moderately actively on the toxic strain, whereasA.bifilosa totally avoided feeding on both strains. This suggeststhat A.bifilosa rejected cyanobacterial filaments due to theirnutritional inadequacy or difficult manageability. The differentresponse of E.affinis to the non-toxic and toxic strains, inturn, shows that this copepod species was able to sense thepresence of the toxin in cyanobacterial filaments and thereforefed less on the toxic strain. The interaction between A.bifilosaand Nodularia sp. was further examined (with the particle countingmethod) by measuring the clearance rates of A.bifilosa on ediblegreen flagellates in the presence of cyanobacteria. The presenceor concentration of toxic Nodularia sp. did not affect grazingrates of A.bifilosa on Brachiomonas submarina. Since earlierstudies have shown that ingestion of Nodularia sp. decreasesegg production and increases mortality in E.affinis, we suggestthat the occurrence of Nodularia sp. blooms in the Baltic Seamay favour individuals of copepod species capable of selectivefeeding, such as A.bifilosa.     

Resource competition between two rotifer species (Brachionus rubens and B.calyciflorus): an experimental test of a mechanistic model

A mechanistic model of competition on a single resource wastested experimentally with two freshwater rotifers, Brachionusrubens and B.calyciflorus, both grown on the alga Chlorellapyrenoidosa. Using open culture systems for each species wemeasured: (i) the resource-saturated exponential growth rate,µ_max, and (ii) the relationship between specific growthrate, µ, at steady-state and the residual algal concentrationover a range of system turnover rates, or dilution rates, D.The µ_max of B.calyciflorus was {small tilde}60% higherthan B. rubens. These results were then used to construct agraphical model for predicting the victor in interspecific competitionbetween the two rotifers. Since the two resource-dependent growthrates crossed, one species, B.calyciflorus, was predicted tobe the victor at a high D while B. rubens was predicted as thevictor at low D. Finally, the outcome of competition was determinedfor two turnover rates. As predicted by the graphical competitionmodels, B.calyciflorus was the dominant species at rapid D (0.029h^–1) and B.rubens was dominant at slow D (0.0044 h^–1).These studies support recent conclusions that mechanistic competitionmodels may be applied to predict dominant species from a prioriinformation on growth potential and resource levels, which isnot possible with traditional Lotka-Volterra models.     

Formation of Radioactive Citrulline during Photosynthetic C14O2-Fixation by Blue-Green Algae

Citrulline has been isolated and identified from extracts ofNostoc muscorum. All members of the Cyanophyceae hitherto investigatedshow a relatively large amount of the CO₂ fixed during photosynthesisin citrulline (ranging as high as 20 per cent. in Nostoc) whencompared to the trace amounts found in the Chlorophyceae. Nostocalso has the ability to fix C¹⁴ in citrulline during dark fixation,but at a rate slower than in light. As no free urea or arginine was found in Nostoc, it is likelythat citrulline is functioning in reactions other than thoseleading to arginine and urea synthesis. Other possible functions for citrulline are briefly discussed.     

Allelopathy of Baltic Sea cyanobacteria: no evidence for the role of nodularin

Extracts of Aphanizomenon flos-aquae and Nodularia spumigena,the two most common cyanobacteria forming recurrent blooms inthe Baltic Sea, decrease the abundance of some phytoplanktonspecies via the release of allelopathic substances. We investigatedhow cell-free filtrates of the two cyanobacteria, as well aspurified hepatotoxin nodularin, produced by N. spumigena affectedcell numbers, chlorophyll a content and ¹⁴CO₂ uptake of thecryptophyte Rhodomonas sp. Both cyanobacterial filtrates significantlyretarded the growth of Rhodomonas sp., A. flos-aquae filtrateup to 46%, whereas purified nodularin showed no significanteffect on any of the growth parameters of the cryptophyte. Theseresults suggest that the allelopathic effect of N. spumigenais most probably due to metabolite(s) other than nodularin,possibly acting via the damage of the target cells.