Publication Date



Open access

Degree Type


Degree Name

Doctor of Philosophy (PHD)


Marine Biology and Fisheries (Marine)

Date of Defense


First Committee Member

Martin Grosell

Second Committee Member

Christopher M. Wood

Third Committee Member

David Letson

Fourth Committee Member

Christopher Langdon

Fifth Committee Member

Lynne Fieber


Freshwater organisms are known to maintain hyperosmotic internal conditions despite outward diffusive loss of ions. The freshwater common pond snail Lymnaea stagnalis faces this challenge while additionally attaining the necessary ions for calcification. These are the first documented assessments of the time and mode of recovery for ions lost due to full-body withdrawal in adults of this species. Additionally, this document reports on the physiological and developmental onset of embryonic calcification and the commencement of active acquisition of shell-forming ions from the surrounding environment. The effect of water chemistry and lead (Pb) exposure on embryonic growth, development, and calcium (Ca2+) acquisition was also tested. Pharmacological and water chemistry manipulations were used to determine mechanisms for embryonic Ca2+ and HCO3-/CO32- acquisition and the sensitivity of those pathways. Lastly, L. stagnalis, was shown to have a lowest effective concentration of -1 using net Ca2+ uptake, growth, and developmental endpoints in laboratory and natural waters. This is the lowest effective concentration observed for any organism to date. One of the most insightful findings reported here is the interconnectedness of the pathways for acquisition of Na+ and Ca2+ through endogenous production of H+ and HCO3- via carbonic anhydrase-catalyzed hydration of metabolic CO2. The combination of high demand for Ca2+ throughout early life stages and periodic acute demands for Na+ recovery following extracellular fluid loss apparently causes L. stagnalis to be highly sensitive to changes in water chemistry, including [Pb] in the embryos, and possibly pH. The findings reported here warn of the need to establish freshwater environmental indicators and consider raising awareness of the threat of freshwater acidification, which may be greater than that of ocean acidification.


Homeostasis; Calcium; Water Chemistry; Invertebrate; Metal Toxicology; Freshwater Physiology