Nutritional deficiencies are a very common phenomenon, and consumers generally face food that is not optimally suited for their needs. Especially herbivores are habitually confronted with food of inferior quality, usually a result of too‐low nutrient concentrations in plant material. Waterfleas of the genus Daphnia are good model organisms to study the effect of inferior quality food, and how animals deal with this. We tested the effect of algae to which we had given different phosphorus contents on both life history and feeding parameters of Daphnia magna. Phosphorus content of the algae strongly affected both the growth rate and the feeding activity of the daphniids. Feeding activity increased with declining food quality (increase in C:P ratio of the algae), whereas growth rates were maximal at intermediate C:P levels. We conclude that the direct limitation of phosphorus is a very important factor determining food quality for zooplankters. Daphniids counterbalanced lower P content of their food by spending more C (energy) on acquiring this limiting resource. This implies that when Daphnia are given phosphorus‐limited food both the addition of phosphorus and the addition of carbon (energy) should increase the growth rate of the animals (co‐limitation). The influence of the phosphorus content of the food on the feeding activity of Daphnia offers a mechanistic explanation for the observed homeostasis in daphniids.
Nutritional imbalances are of great interest in the ecological stoichiometry literature, in which researchers have focused almost exclusively on cases where nutrients are available in low amounts relative to energy (carbon), and animal growth is impaired due to insufficient nutrient intake. Little attention has been given to situations where food elemental content is higher than the level that satisfies animal requirements. However, most animals are strongly homeostatic with respect to the elemental composition of their body; hence they must excrete the excess of elements that are not in short supply. To date, stoichiometric theory has assumed that excretion of superfluous elements does not come with a cost and, thus, that consumption of food with surplus nutrients does not impair performance. Here we challenge this assumption, based on a compilation of several examples involving food phosphorus content that show that the performance of a wide variety of animals decreases when supplied with food containing high concentrations of (potentially) limiting nutrients. We discuss possible mechanisms for this phenomenon, and suggest that animals most vulnerable to effects of high food nutrient content are those that normally feed on low- quality (low-nutrient: C) food, and have a relatively low body nutrient content themselves, such as herbivores and detritivores.
Analyzing one of the most extensive long-term data series in the North Sea, the Helgoland Roads time series, we investigated the changes in the factors that potentially drive phytoplankton bloom dynamics in the German Bight. We compared the changes in these factors with the changes in the spring bloom phenology. We combined zooplankton, nutrient, weather, and phytoplankton data to analyze whether there has been a shift in trophic interactions in the North Sea affecting the spring bloom timing. The potential influence of temperature, with a mean increase of 1.5uC, was investigated. We showed that the German Bight around Helgoland is a highly dynamic system and has undergone considerable change in the last 30 yr. Nutrient levels, temperature, underwater light climate and wind speed have all changed. However, the spring bloom dynamics have hardly changed at all. We showed that the spring bloom tends to come later in warmer years but that this is not directly correlated with the overall warming trend. The known regime shift of the late 1980s is clearly visible in our data in terms of average phytoplankton winter densities and average cell size, but even so the start of the spring bloom has not changed.
An experiment was conducted to measure DE and ME and the apparent total tract digestibility (ATTD) of energy, N, and P in distillers dried grains with solubles (DDGS) fed to growing pigs. Ten sources of DDGS were obtained from ethanol plants in South Dakota and Minnesota, and 11 diets were formulated. One diet was based on corn (96.8%), limestone, salt, vitamins, and microminerals. Ten additional diets were formulated by mixing the corn diet and each of the 10 sources of DDGS in a 1:1 ratio. Eleven growing pigs (initial BW of 29.3 +/- 0.42 kg) were allotted to an 11 x 11 Latin square design, with 11 periods and 11 pigs. Each of the 11 diets was fed to each pig during 1 period. Pigs were placed in metabolism cages that allowed for the total, but separate, collection of feces and urine. Samples were analyzed for GE, N, and P and energy and N balances, and the ATTD of GE, N, and P were calculated for each diet. By subtracting the contribution from the corn diet to the DDGS-containing diets, the energy and N balances and the ATTD for GE, N, and P for each source of DDGS were calculated. Results of the experiment showed that the DE and ME differed (P < 0.001) among the 10 sources of DDGS (3,947 to 4,593 kcal of DE/kg of DM and 3,674 to 4,336 kcal of ME/kg of DM). The average DE and ME in DDGS were 4,140 and 3,897 kcal/kg of DM, respectively. These values were not different from the DE and ME in corn (4,088 and 3,989 kcal/kg of DM, respectively). Based on the analyzed GE and nutrient composition of DDGS and the calculated values for DE and ME, prediction equations for DE and ME were developed. These equations showed that DE and ME in DDGS may be predicted from the concentration of ash, ether extract, ADF, and GE. The retention of N from DDGS was greater (P < 0.001) than from corn, but when calculated on a percentage basis, the N retention did not differ between DDGS and corn. The ATTD of P in DDGS was 59.1% on average for the 10 samples. This value was greater (P < 0.001) than the ATTD of P in corn (19.3%). It is concluded that the DE and ME in DDGS is not different from the DE and ME in corn. However, if DDGS is included in diets fed to growing swine, a greater portion of the organic P will be digested and absorbed, thus reducing the need for adding inorganic P to the diets.
The objective of this experiment was to measure the digestibilities of energy, CP, and AA in 10 samples of corn distillers dried grain with solubles (DDGS) and in corn fed to growing pigs. Twelve growing barrows (initial BW: 34.0 +/- 1.41 kg) were allotted to an 8 x 12 Youden square design with 8 periods and 12 animals. Ten of 12 diets were based on the 10 DDGS samples (66.7%), 1 diet was based on corn (97%), and the last diet was a N-free diet based on cornstarch and sucrose. Chromic oxide (0.3%) was included in all diets as an inert marker. Pigs were provided their respective diets at a level of 3 times their estimated energy requirement for maintenance. The apparent (AID) and standardized (SID) ileal digestibilities for CP and AA were measured in the 10 samples of DDGS and in corn using the direct procedure, but the apparent total tract digestibilities for DM and GE were estimated using the difference procedure. The concentration of DE in each sample of DDGS and in corn was also calculated. The results of the experiment indicated variation among the different sources of DDGS in AID and SID for Lys, which ranged from 35.0 to 55.9% and 43.9 to 63.0%, respectively. For Met, the SID varied between 73.9 and 84.7%. However, the variability among samples in the SID for CP, and for the indispensable AA other than Lys and Met, was relatively low and ranged between 6 and 8 percentage units (i.e., from 64.0 to 70.6%, 74.1 to 80.1%, and 67.4 to 75.3% for Thr, Trp, and Ile, respectively). The SID for Trp in corn (72.8%) was lower (P < 0.05) than in DDGS, but for the remaining indispensable AA, except Arg, the SID for corn were greater (P < 0.01) than for DDGS. The DE concentration in the 10 samples of DDGS varied (P < 0.001) from 3,382 to 3,811 kcal of DE per kg of DM. For corn, the DE was 3,845 kcal per kg of DM. It is concluded that the AID and SID for Lys vary among samples of DDGS, but for most other AA the AID and SID are relatively similar and vary only 6 to 8 percentage units among different samples. Future work should focus on identifying the reasons for the variation in the digestibility of Lys to avoid processing procedures that are detrimental to Lys digestibility.
We investigated whether nutrient limitations of primary producers act upward through food webs only in terms of density effects or if there is a second pathway for nutrient limitation signals channelled upward to higher trophic levels. We used tritrophic food chains to assess the effects of nutrient-limited phytoplankters (the cryptophyte Rhodomonas salina) on herbivorous zooplankters (the calanoid copepod Acartia tonsa) and finally zooplanktivores (larval herring Clupea harengus) living on the herbivores. The primary producers' food quality had a significant effect on fish condition. Our experimental phosphorus-limited food chain resulted in larval fish with a significantly poorer condition than their counterparts reared under nitrogen-limited or nutrient-sufficient conditions. Our results show that mineral nutrient requirements of consumers have to be satisfied first before fatty acids can promote further growth. This challenges the match/mismatch hypothesis, which links larval fish survival probability solely to prey availability, and could imply that reduced nutrient releases into the environment may affect fish stocks even more severely than previously believed.
We investigated the effects of various mineral and biochemical limitations on Daphnia magna. These daphniids have much lower saturation thresholds for growth for the polyunsaturated fatty acids (PUFA) eicosapentaenoic acid (EPA), and arachidonic acid (ARA) than has been previously described for other Daphnia species. Daphniids take up large amount of fatty acids from food, and different fatty acids are handled differently by D. magna. The saturated fatty acid (20:0; EPA) was not retained, and metabolized, the PUFAs were preferably stored. There were also differences among the PUFAs: EPA was found in higher concentrations in the eggs than ARA. In contrast, although there were some variations in D. magna phosphorus levels with varying levels of phosphorus in the food, these differences were small compared with the changes in D. magna fatty-acid concentrations. Independent of these small changes, the P content of eggs was constant at 14 mg P (g dry wt)Ϫ1 . Storage of EPA, but not P, fully compensated D. magna growth during periods of bad food quality. Egg production was a major drain of fatty acids from female D. magna.
In their seminal paper, Goldman et al. suggested that phytoplankton close to maximum growth rate attains a restricted optimal N : P ratio close to the Redfield ratio of molar N : P = 16. Recently, the presence of such a global attractor for optimal phytoplankton stoichiometry has been questioned in models and empirical analyses. As the chemical composition of phytoplankton is of major importance for our understanding of global elemental cycles and biogeochemical transformations, we assembled 55 data sets of phytoplankton growth rate and biomass N : P ratios in a meta‐analysis testing (1) whether phytoplankton N : P converges at high growth rates, (2) whether N : P ratios scale with growth rate, and (3) whether the optimal N : P ratios achieved at highest growth rates reflect organism traits or environmental conditions. Across systems and species, phytoplankton N : P decreased with increasing growth rate and at the same time showed decreasing variance, i.e., fast‐growing phytoplankton is more P rich and has a more confined elemental composition. Optimal N : P increased with increasing N : P of available nutrients, i.e., with increasing P limitation. Other differences were rare, except cyanobacteria showed higher optimal N : P than diatoms. Understanding the role of phytoplankton in biogeochemical transformation requires modeling approaches that are stoichiometrically flexible to reflect the dynamics of growth and nutrient supply in primary producers.
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