The mean carbon, nitrogen, and phosphorus contents of particulate material for 5 1 lakes or lake basins, extending from arctic to tropical climatic regions, including small lakes as well as the largest lakes in the world, indicate that Redfield ratios are the exception rather than the rule in freshwater. The C: P and N : P ratios are more variable for lake particles but generally higher than marine particles, and the mean molar C: N, C: P, and N : P ratios are substantially higher than the Redfield ratio of 106 : 16: 1. On average, lower C : N, C : P, and N : P ratios occur in subarctic lakes while higher ratios occur in the tropics and in temperate, oligotrophic lakes on the Canadian Shield. In shield lakes with long residence times (>6 months) the high ratios of C : N, C : P, and N : P do not originate from streamborne or atmospherically deposited particles but arise from in-lake processes. Regression analysis demonstrates that small lakes are generally more N and P deficient than large lakes. In freshwaters, particulate composition ratios imply that a wide variety of conditions exists in lakes, including N and P deficiency, as well as N and P sufficiency. In the Experimental Lakes Area of Canada, independent physiological nutrient status indicators generally agree with the status indicated by seston ratios. The relative uniformity of marine C : N : P composition (compared to lakes) at the Redfield ratio suggests that marine plankton cannot be as severely, or as frequently, limited by N and P as lake plankton. Consequently, the paradigm of N limitation in the oceans requires qualification.Based on particulate comDosition, it is more correct to say that ocean plankton is noi as N grid P deficient as lake-plankton.The composition of marine particulate matter is relatively uniform. Redfield (1934Redfield ( , 1958 noted the near constancy of the ratio of C : N : P in marine plankton and the similarity of the N : P ratio of plankton to the oceanic deepwater ratio of nitrate to phosphate. As early as 1940, the C : N : P molar composition ratio of marine plankton was accepted to be 106 : 16 : 1 (Redfield et al. 1963); this ratio is now referred to as the Redfield ratio. The ratio has withstood the test of time, and the ever-growing number of analyses of marine particles and nutrient regeneration profiles, with relatively AcknowledgmentsWe thank G. J. Bumskill and H. E. Welch for permission to publish particulate C, N, and P data on Lake Winnipeg and the Saqvaqjuac lakes, respectively. Dana Cruikshank and his field assistants sampled the ELA lakes for us.

1. Within a lake district of relatively homogeneous geomorphology, the responses of lakes to climate are influenced by the complexity of the hydrogeologic setting, position in the landscape, and lake‐specific biological and physical features. We examined lake chemical responses to drought in surface water‐ and groundwater‐dominated districts to address two general questions. (1) Are spatial patterns in chemical dynamics among lakes uniform and synchronous within a lake district, suggesting broad geomorphic controls; variable in a spatially explicit pattern, with synchrony related to landscape position, suggesting hydrologic flowpath controls; or spatially unstructured and asynchronous, suggesting overriding control by lake‐specific factors? (2) Are lake responses to drought a simple function of precipitation quantity or are they dictated by more complex interactions among climate, unique lake features, and hydrologic setting? 2. Annual open‐water means for epilimnetic concentrations of chloride, calcium, sulfate, ANC, DOC, total nitrogen, silica, total phosphorus, and chlorophyll a measured between 1982 and 1995 were assembled for lakes in the Red Lake and ELA districts of north‐western Ontario, the Muskoka – Dorset district in south‐central Ontario, and the Northern Highland district of Wisconsin. Within each district, we compared responses of lakes classified by landscape position into highland or lowland, depending on relative location within the local to regional hydrologic flow system. Synchrony, defined as a measure of the similarity in inter‐annual dynamics among lakes within a district, was quantified as the Pearson product‐moment correlation (r) between two lakes with observations paired by year. To determine if solute concentrations were directly related to interannual variations in precipitation quantity, we used regression analysis to fit district‐wide slopes describing the relationship between each chemical variable and annual (June to May) and October to May (Oct–May) precipitation. 3. Among lakes in each of the three Ontario districts, the pattern of chemical response to interannual shifts in precipitation was spatially uniform. In these surface water‐ dominated districts, solute concentrations were generally a simple function of precipitation. Conservative solutes, like calcium and chloride, tended to be more synchronous and were negatively related to precipitation. Solutes such as silica, total phosphorus, and chlorophyll a, which are influenced by in‐lake processes, were less synchronous and relationships with precipitation tended to be positive or absent. 4. In the groundwater‐dominated Northern Highland lakes of Wisconsin, we observed spatial structure in drought response, with lowland lakes more synchronous than highland lakes. However, there was no evidence for a direct relationship between any solute and precipitation. Instead, increases in the concentration of the conservative ion calcium during drought were not followed by a symmetrical return to pre‐drought conditions when precipitation ...

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Results of a 4-yr study of the hydrology of the Rawson Lake watershed, as well as the chemistry of precipitation, streamflow, and the lake are presented. Inputs of water and most chemicals varied by as much as 2 times from one year to the next, illustrating that several years’ data are necessary to determine accurately average nutrient inputs and losses from an ecosystem. Precipitation appeared to be almost the sole source of P and N to lake and terrestrial ecosystems, although N2 fixation cannot be dismissed in the latter case. Significant proportions of Ca, Mg, Na, K, Cl, and SO4 were supplied directly from precipitation. The lake also received substantial quantities of these elements as well as silicon, from weathering of terrestrial geological materials. Although far from any major industrial sources of SO2, the pH of precipitation averaged < 5 for the years studied. Both terrestrial and lake systems retained a high proportion of entering P and N, and smaller proportions of other nutrients. Interpretation of relative retentions of nutrients revealed that the terrestrial ecosystem has well-balanced P and N supplies, while the lake was P limited. Rates of loss of chemicals from the terrestrial watershed of Rawson Lake are comparable to other sites in the Precambrian Shield, but lower than values from other geological settings.

The ability of nutrients to control photosynthesis was compared in epilithon (the association on rock surfaces in the littoral zone) and phytoplankton of 13 low alkalinity lakes of the Experimental Lakes Area of northwestern Ontario. The study included (1) surveys of lakes varying in nutrient concentrations; (2) experimental additions to lakes of carbon and nitrogen (N), with or without phosphorus (P); and (3) experimental additions to lakes of sulfuric and nitric acids. Nutrient controls of planktonic and epilithic algal photosynthesis differed consistently. Phosphorus limited planktonic algal photosynthesis. In contrast, dissolved inorganic carbon (DIC) limited epilithic photosynthesis in both perturbed and unperturbed lakes because diffusive resistance kept the effective supply of DIC below the level needed for optimal growth. Epilithic photosynthesis was lowered when lake disturbances (e.g., acidification) reduced epilimnetic concentrations of DIC. Expected increases in atmospheric carbon dioxide can, therefore, differentially affect the littoral and pelagic food webs in low DIC lakes. Epilithic photosynthesis in all study lakes was unrelated to N or P availability despite apparent N and P deficiencies, based upon particulate nutrient ratios. Rates of epilithic respiration were, however, correlated with epilimnetic concentrations of inorganic N.