Grazer species effects on epilithon nutrient composition

1. Field and laboratory experiments were conducted to investigate the excretion stoichiometry of nitrogen (N) and phosphorus (P) of two benthic macroinvertebrate grazers, the crayfish Orconectes propinquus and the snail Elimia livescens, that differ in body stoichiometry (mean body molar N : P 18 and 28, respectively). Crayfish excretion had a significantly higher ammonium : soluble reactive phosphorus (SRP) ratio in the laboratory and in three natural streams than did snails, as predicted by ecological stoichiometry theory. 2. In greenhouse recirculating artificial streams, treatments consisting of crayfish, snails, or no grazers were used to examine responses in dissolved nutrient concentrations and epilithon nutrient composition and limitation. SRP concentrations depended upon the grazer species, with the snail treatment having a higher SRP concentration than other treatments (P < 0.05). Dissolved inorganic N was not affected by grazers, but appeared to be rapidly incorporated in epilithon. 3. Epilithon N content was dependent upon the grazer species present, with the crayfish treatment having a significantly higher N content than other treatments (P = 0.001). No grazer species effects on epilithon P content were found. However, both grazer treatments had significantly lower epilithon P content than the no‐grazer treatment. 4. Traditionally, studies have focused on how grazer‐induced structural changes to epilithon can alter epilithon nutrient dynamics, but this structural mechanism could not solely explain differences in epilithon nutrient contents and ratios in the present study. Our results rather suggest that benthic grazers can alter epilithon nutrient composition and limitation via nutrient excretion. Consequently, macroinvertebrate grazers may serve as ‘nutrient pumps’ that partly regulate the availability of nutrients to algae in stream ecosystems.     

Crayfish predation on the common pond snail (Lymnaea stagnalis): the effect of habitat complexity and snail size on foraging efficiency

Optimal foraging theory was used to explain selective foraging by the introduced signal crayfish (Pacifastacus leniusculus) on the thin-shelled common pond snail (Lymnaea stagnalis). Crayfish predation efficiency was studied in relation to habitat complexity and snail size. In a pool experiment (area 1.3 m²) single adult crayfish were allowed to feed on four size classes of snails for one week. A pair-wise preference trial (aquarium experiment) tested if adult crayfish selectively predated on particular size classes of snail and if prey value (expressed as snail dry mass per handling time) could explain the size range of snails chosen. Crayfish preferred the smallest size classes of snails in both pool and aquaria experiments. In the pool experiment crayfish had a strong effect on snail survival. Habitat complexity did not affect overall snail survival, but resulted in reduced predation pressure on the smallest size classes of snails. Handling time and shell-thickness increased exponentially with increasing snail size, and the two smallest size classes had the highest prey values. The results suggest that crayfish can structure the abundance and size distribution of thin-shelled snails, through size-selective predation and reduction of macrophytes. The mechanisms behind the choice of snails may be based on prey value and reduced exposure time to predators and conspecifics. Crayfish effects on snail size distribution may be less pronounced in complex habitats such as macrophyte beds. This revised version was published online in July 2006 with corrections to the Cover Date.     

The independent and interactive effects of snail grazing and nutrient enrichment on structuring periphyton communities

We investigated the independent and interactive effects of nutrient enrichment and snail grazing on structuring periphyton communities in a northern temperate lake. Nutrient releasing substrates and grazer enclosures were used to simultaneously manipulate nutrient availability and herbivory. Periphyton was allowed 18 days to accrue before grazers (Elimia livescens = Goniobasis livescens) were introduced.Addition of nitrogen and phosphorus caused a significant increase in biovolume (p < 0.001), whereas grazing had no significant effect on biovolume but resulted in a shift in species composition. Four taxa were largely responsible for the increase in biovolume on the nutrient enriched substrates: Oedogonium sp, Stigeoclonium tenue, Navicula radiosa var. radiosa and Navicula radiosa var. tenella. By the 28th day, nutrient enrichment caused a shift from a community dominated by diatoms (Bacillariophyceae) to a community dominated by green algae (Chlorophyceae). Blue green algae (Myxophyceae) maintained an equal proportion in high and low-nutrient regimes.Grazing had a more pronounced effect on altering community composition on the nutrient enriched substrates than on the unenriched substrates. Grazing caused a decrease in diversity and an increase in dominance by green algae on the nutrient enriched substrates. The relative biovolume of green algae increased from 64% to 93% on grazed substrates, due to the significant increase in relative abundance of Stigeoclonium tenue. This taxon has both prostate basal cells and erect filamentous cells. The ratio of basal: filamentous cells increased from 4.7 to 5.2 with grazing, suggesting that the heretotrichous growth form of Stigeoclonium tenue is adapted to grazing by virtue of the basal cells which are able to adhere to the substratum and resist being grazed.     

Ammonia Excretion in Snails Viviparus bengalensis (LAMARCK) and Pila globosa (SWAINSON) during Active and Dormant Periods

The quantities of ammonia excreted over a period of 24 h by two prosobranch snails Viviparus bengalensis and Pila globosa were measured at 30 day intervals to compare their excretion during the active and dormant phases of their life. The aquatic snail V. bengalensis excreted 0.5-1.0 μmole of ammonia per day and snail throughout the year. The ammonia excretion rate of this snail was 40-50% lower in winter than in the monsoon and summer period. In the amphibious snail Pila globosa excretion of ammonia was limited to only four months of its active life i. e. during July-October. Also the rate of ammonia excretion in P. globosa (0.2-0.3 μmole per day and snail) was very low compared with V. bengalensis.     

The diatoms ingested by freshwater snails: temporal,spatial, and interspecific variation

Seventeen species of diatoms, representing a broad range of sizes, shapes, and growth habits, were collected from rocks in rapidly-flowing sections of the Mitchell River, North Carolina. The diatoms ingested by adult Goniobasis proxima, juvenile Leptoxis carinata, and adult Physa sp. co-occurring in this habitat were indistinguishable from one another, in spite of great differences in radular morphology. All snails sampled the diatom flora almost randomly, with only one or two of the larger diatom species under-represented in the gut contents. Some snails also seemed to selectively ingest the smaller individuals of the larger diatom taxa, and larger individuals of the smaller diatom taxa. The diatoms identifiable in juvenile Goniobasis guts were somewhat more distinctive, although this seemed to be due at least partly to more mechanical breakage. The diatom flora of quiet, muddy pools was much different from that of shallow, rocky areas, but once again, Goniobasis seemed to sample the available flora randomly. Seasonal variation was also apparent in the diatom diet of Goniobasis. We suggest that in some cases, it may be reasonable to use snails to sample the diatom assemblage present in a particular habitat, if more direct methods are impractical.     

Cell characteristics and biochemical composition ofDunaliella primolecta Butcher conditioned at different concentrations of dissolved nitrogen

The rates of growth, cell size, elemental and biochemical composition ofDunaliella primolecta were monitored during exponential growth after conditioning over three weeks in media containing different concentrations of nitrogen. The rate of growth, measured both as cell density and cell volume, was correlated positively with the N concentration of the medium (P<0.01). N-starved cells did not grow and died within three days. Cell volume and dry weight per cell were significantly higher (P<0.01) in the N-low (0.45 g-at 1^–1 NO₃-N) condition than in the N-high I condition (3.53 g-at 1^–1 NO₃-N). In the N-high II condition the addition of 1.87 g-at 1^–1 NH₄-N to 3.53 g-at 1^–1 NO₃-N did not significantly enhance growth.The composition was influenced by the availability of nitrogen. The amount of protein per cell decreased significantly from 20.1 to 9.7 pg with reduced nitrate availability between the N-high condition and the N-low condition (P<0.01). The associated carbohydrate content increased significantly from 10.6 to 26.74 pg per cell (P<0.001). These findings confirm the potential for significant variability in the composition ofD. primolecta, both in the natural environment and following experimental manipulation.     

Spatial and seasonal variation in nutrient excretion by benthic invertebrates in a eutrophic reservoir

1. Nitrogen (N) and phosphorus (P) fluxes via excretion by benthic invertebrates were quantified in a eutrophic reservoir (Acton Lake, Ohio, U.S.A.). We quantified variation in nutrient fluxes seasonally (June until November 1997), spatially (three sites) and among taxa (chironomids, tubificid oligochaetes and Chaoborus). 2. The three taxa differed in spatial distribution and contribution to nutrient fluxes. Tubificids were the most abundant taxon at two oxic sites (1.5 and 4 m depth), and were exceedingly rare at an anoxic, hypolimnetic site (8 m). Chironomids were abundant only at the shallowest oxic site. Chaoborus was the only abundant taxon at the anoxic site. Total benthic invertebrate biomass was greatest at the shallowest site and lowest at the anoxic, hypolimnetic site. 3. Mass‐specific excretion rate [μmol NH₄–N or soluble reactive P (SRP) excreted mg dry mass^–1 h^–1] varied among experiments and was influenced by temperature. Differences among taxa were not significant. Thus, nutrient flux through benthic invertebrates was affected more by total invertebrate biomass and temperature than by species composition. 4. Fluxes of N and P via benthic invertebrate excretion (μmol NH₄–N or SRP m^–2 day^–1) were greatest at the oxic sites, where fluxes were dominated by the excretion of tubificids and chironomids. The N and P fluxes at the anoxic site were much lower, and were dominated by excretion by Chaoborus. The ratio of N and P excreted by the benthic invertebrate assemblage varied seasonally and was lowest at the anoxic site. 5. Comparison with other measured inputs shows that excretion by benthic invertebrates could be an important source of nutrients, especially of P. However, the relative importance of nutrient excretion by the benthos varies greatly spatially and temporally.     

Signatures of nitrogen limitation in the elemental composition of the proteins involved in the metabolic apparatus

Nitrogen (N) is a fundamental component of nucleotides and amino acids and is often a limiting nutrient in natural ecosystems. Thus, study of the N content of biomolecules may establish important connections between ecology and genomics. However, while significant differences in the elemental composition of whole organisms are well documented, how the flux of nutrients in the cell has shaped the evolution of different cellular processes remains poorly understood. By examining the elemental composition of major functional classes of proteins in four multicellular eukaryotic model organisms, we find that the catabolic machinery shows substantially lower N content than the anabolic machinery and the rest of the proteome. This pattern suggests that ecological selection for N conservation specifically targets cellular components that are highly expressed in response to nutrient limitation. We propose that the RNA component of the anabolic machineries is the mechanistic force driving the elemental imbalance we found, and that RNA functions as an intracellular nutrient reservoir that is degraded and recycled during starvation periods. A comparison of the elemental composition of the anabolic and catabolic machineries in species that have experienced different levels of N limitation in their evolutionary history (animals versus plants) suggests that selection for N conservation has preferentially targeted the catabolic machineries of plants, resulting in a lower N content of the proteins involved in their catabolic processes. These findings link the composition of major cellular components to the environmental factors that trigger the activation of those components, suggesting that resource availability has constrained the atomic composition and the molecular architecture of the biotic processes that enable cells to respond to reduced nutrient availability.     

The chemical composition of Astragalus: a comparison of seleniferous and non-seleniferous plants growing side by side

COWGILL, U. M. & LANDENBERGER, B.D., 1992. The chemical composition of Astragalus: a comparison of seleniferous and non-seleniferous plants growing side by side. This paper describes how non-seleniferous plant species coexist with seleniferous ones and what chemical changes occur in non-seleniferous species that allow the toleration of large quantities of volatilized Se-bearing compounds. These compounds are known for their phytotoxicity as well as for their toxicity to mammals and insects. Twenty-three sites (Colorado, Utah, New Mexico, U.S.A.) were examined over a 6-year period. Plants collected from these sites were divided into four groups: seleniferous astragali, seleniferous Brassicaceae, non-seleniferous astragali and non-seleniferous associated genera. Furthermore, it was possible to categorize these sites: Type 1 sites supported all four groups of plants, Type 2 sites contained only seleniferous astragali, whereas Type 3 sites sustained only non-seleniferous astragali. When concentrations of Cu, Zn (P <0.0001), Si (P<0.02), As, Pb (P<0.03) and Cs (P<0.05) were measured, the values for those seleniferous astragali that coexisted with other species were significantly different from the values found for those seleniferous astragali that grew alone. There were four areas that in some years supported all four groups of plants (Type 1 site) and in other years contained only seleniferous astragali. In this way, the latter may be used as a chemical control for the former. The seleniferous astragali collected at Type 1 sites always contained more Cu, As, Si and Pb than the solitary astragali of Type 2 sites. When seleniferous astragali of Type 1 sites that support all four groups of plants develop in a year where the usually associated species of past years are absent, they contain in their tissues concentrations of Cu, As, Si, Pb, Zn and Cs typical of Type 2 sites where the seleniferous astragali grew alone. Non-seleniferous astragali of Type 1 sites that support all four groups of plants have a chemical composition that differs significantly in the quantity of 15 elements from non-seleniferous astragali of Type 3 sites where the latter grew alone. Since allelochemicals such as phenolic acids and flavonoids are well known to be able to alter mineral absorption by plants and since the astragali are known to produce such substances, it is suggested that the possible production of phenolic acids and flavonoids may permit coexistence of seleniferous plants with non-seleniferous ones and thus explain the chemical differences among the plants of the four types of sites.     

放牧对五台山高山、亚高山草甸牧草品质的影响

通过野外样地调查和室内分析,从牧草的适口性和营养成分两方面,研究了放牧对五台山高山、亚高山草甸牧草品质的影响。结果表明：放牧干扰草甸中适口性为喜食及以上的优质牧草种类为20种,喜食牧草在群落中的相对盖度最大,为45.40%;与之相比,极度退化草甸中优质牧草种类为13种,乐食牧草的相对盖度最大,为62.24%;喜食以上优质牧草的相对盖度最大值,出现在轻度退化草甸中,达到65.95%。同一适口性等级牧草,在不同退化等级草甸中,种类发生变化。从放牧干扰草甸到极度退化草甸,牧草中的粗蛋白和灰分含量呈增加趋势,而中性洗涤纤维和酸性洗涤纤维含量呈下降趋势。放牧压力下,五台山高山、亚高山草甸的牧草品质并未下降,但放牧造成的草甸生产力降低,制约了资源的可持续利用和保护。     

Whole body elemental and proximate composition of Atlantic salmon (Salmo salar) during the life cycle

Atlantic salmon ( Salmo salar ) raised under production conditions were sampled periodically during their life cycle, in fresh and salt water, to determine their proximate and elemental composition. The parameters changed in relation to life cycle stage and fish size, and were influenced by whether the fish were in salt or freshwater. Measurements presented can be used to assist in the formulation of feeds, and to determine the health and nutritional status of commercially reared Atlantic salmon.     

高摄食压力下球形棕囊藻凝聚体的形成

棕囊藻包含囊体和游离单细胞两种生活史阶段。囊体是棕囊藻藻华爆发时的优势形态,藻华衰退时囊体能够形成凝聚体,但是棕囊藻游离单细胞的凝聚体极少被发现。本次研究将球形棕囊藻单细胞和高密度的海洋弯曲甲藻共同培养,使球形棕囊藻的生长承受高摄食压力,通过观察摄食者和棕囊藻的生长、凝聚体的数量和形态,阐明单细胞凝聚体的形成以及与摄食压力的关系。当球形棕囊藻进入衰退期时,高摄食压力引发游离单细胞聚合形成凝聚体,无摄食压力情况下,单细胞不形成凝聚体。凝聚体由无鞭毛细胞组成,细胞排列紧密,近似球体。凝聚体形成伊始,凝聚体内部可见凝胶状物质将细胞互相粘结,并且粘附了纤维等物质。凝聚体的体积和粘附的细胞数量逐渐提高,细胞排列愈加紧密,凝聚体内部形态和结构不易分辨。凝聚体的形成有效保护了部分单细胞免受摄食压力的影响,减少了摄食死亡率。凝聚体的形成是球形棕囊藻面临高摄食压力时采取的主动的防御策略。球形棕囊藻能够频繁引发大规模藻华的原因可能在于其在生长的各个阶段中均具有优越的竞争策略。     

Effects of phosphorus enrichment and grazing snails on modern stromatolitic microbial communities

1. The effects of phosphorus enrichment and grazing snails on a benthic microbial community that builds stromatolic oncolites were examined in an experiment at Rio Mesquites, Cuatro Ciénegas, Mexico. Chemical analyses of stream water samples indicated that overall atomic ratios of total nitrogen (N) to total phosphorus (P) were approximately 110, indicating a strong potential for P‐limitation of microbial growth. 2. Phosphorus enrichment involved addition of 5 μmol Na₂HPO₄ L^?1 to streamside microcosms receiving intermittent inputs of stream water while grazer manipulation involved removal of the dominant grazer, the snail Mexithauma quadripaludium. After 7 weeks, we examined responses in organic matter content, C : N : P ratios, metabolism (P removal, primary production, dark respiration, and calcification), and microbial community structure using molecular fingerprinting of 16S rRNA genes. 3. Manipulation of snails did not affect response variables measured in these treatments (organic matter, C : P ratio, P removal rate). However, P enrichment significantly decreased the C : P and N : P ratios of surficial materials in the oncolites (organic matter content was unchanged), increased net and gross photosynthesis (oxygen consumption in the dark was unchanged), increased rates of calcification, and increased diatoms relative to cyanobacteria. Heterotrophic Eubacteria and Archaea were only modestly affected. Thus, our results indicate weak grazing effects but strong impacts of P in this benthic system. 4. We hypothesise that a state of severe P‐limitation is imposed on autotrophic production in this food web due, at least in part, to co‐precipitation of phosphate during calcite deposition. This produces severe P‐limitation of the benthic algae and cyanobacteria, resulting in high C : P ratio of microbial mats relative to the biomass of photoautotrophs (phytoplankton, terrestrial foliage) in other ecosystems. In turn, this high C : P ratio is likely to generate severe stoichiometric constraints on the herbivores, thus limiting their populations and resulting in weak overall grazing impacts.     

宁夏沙坡头地区藻类及其分布

通过野外采样,室内培养、观察、鉴定,研究了沙坡头地区藻类多样性和分布特点,并为当地藻类资源利用提出建议。研究发现沙坡头地区藻类植物共计40种(包括1变型),其中蓝藻17种,绿藻10种,硅藻9种,裸藻4种,全为普生种,陆地生境中种类最丰富。亚气及陆地生境中蓝藻在种类数(除灌溉林地结皮)、生物量方面占主导地位;贫瘠水体生境中硅藻占主要地位,营养较丰富水体中绿藻处于优势地位。另外,陆生生境中多以具鞘微鞘藻Microcoleus vaginatus为优势种,小席藻Phormidium tenue为主要种。     

EFFECT OF NUTRIENT AVAILABILITY ON THE BIOCHEMICAL AND ELEMENTAL STOICHIOMETRY IN THE FRESHWATER DIATOM STEPHANODISCUS MINUTULUS (BACILLARIOPHYCEAE)*

The objective of this study was to examine the differences in the biochemical and elemental stoichiometry of a freshwater centric diatom, Stephanodiscus minutulus (Grun.), under various nutrient regimes. Stephanodiscus minutulus was grown at μ_max or 22% of μ_max under limitation by silicon, nitrogen, or phosphorus. Cell sizes for nutrient‐limited cultures were significantly smaller than the non‐limited cell sizes, with N‐limited cells being significantly smaller than all other treatments. Compared with the nutrient‐replete treatment, both carbohydrates and lipids increased in Si‐ and P‐limited cells, whereas carbohydrates increased but proteins decreased in N‐limited cells. All of the growth‐limited cells showed an increase of carbohydrate and triglyceride, and a decrease of cell size and polar lipids as a percentage of total lipids. The non‐limited cells also had a significantly higher chl a concentration and galactolipids as a percentage of total lipids than any of the limited treatments, and the low‐Si and low‐P cells had significantly higher values than the low‐N cells. The particulate C concentrations showed significant differences between treatments, with the Si‐ and P‐limited treatments being significantly higher than the N‐ and non‐limited treatments. Particulate Si did not show a strong relationship with any of the parameters measured, and it was the only parameter with no differences between treatments. The low‐Si cells had a significantly higher P content (about two times more) than any other treatment, presumably owing to the luxury consumption of P, and a correspondingly high phospholipid concentration. The elemental data showed that S. minutulus had a high P demand with low optimum N:P (4) and Si:P (10) ratios and a C:N:P ratio of 109:16:2.3. The particulate C showed a positive relationship with POM (r = 0.93), dry weight (r = 0.88), lipid (r = 0.87) and protein (r = 0.84, all P < 0.0001). Particulate N showed a positive relationship with galactolipids (r = 0.95), protein (r = 0.90), dry weight (r = 0.78), lipid (r = 0.75), and cell volume (r = 0.64, all P < 0.0001). It is evident that nutrient limitation in the freshwater diatom S. minutulus has pronounced effects on its biochemical and elemental stoichiometry.