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CORVALLIS, Ore. - The mortality of larval Pacific oysters in Northwest hatcheries has been linked to ocean acidification, yet the rate of increase in anthropogenic carbon dioxide in the atmosphere and the decrease of pH in near-shore waters have been questioned as being severe enough to cause the die-offs.

However, a new study of Pacific oyster and Mediterranean mussel larvae found that the earliest larval stages are directly sensitive to saturation state, not carbon dioxide (CO₂) or pH. Saturation state is a measure of how corrosive seawater is to the calcium carbonate shells made by bivalve larvae, and how easy it is for larvae to produce their shells.

It is important to note that increasing CO₂ lowers saturation state, the researchers say, and saturation state is very sensitive to CO₂; the challenge interpreting previous studies is that saturation state and pH typically vary together with increasing CO₂. The scientists utilized unique chemical manipulations of seawater to identify the direct sensitivity of larval bivalves to saturation state.

Results of the study, which was funded by the National Science Foundation, are being reported this week in the journal Nature Climate Change.

"Bivalves have been around for a long time and have survived different geologic periods of high carbon dioxide levels in marine environments," said George Waldbusser, an Oregon State University marine ecologist and biogeochemist and lead author on the study, "The difference is that in the past, alkalinity levels buffered increases in CO₂, which kept the saturation state higher relative to pH."

"The difference in the present ocean is that the processes that contribute buffering to the ocean cannot keep pace with the rate of anthropogenic CO₂ increase," added Waldbusser, who is in Oregon State's College of Earth, Ocean, and Atmospheric Sciences. "As long as the saturation state is high, the oysters and mussels we tested could tolerate CO₂ concentrations almost 10 times what they are today."

The idea that early bivalve development and growth is not as physiologically linked to CO₂ or pH levels as previously thought initially seems positive. However, the reverse is actually true, Waldbusser noted. Larval oysters and mussels are so sensitive to the saturation state (which is lowered by increasing CO₂) that the threshold for danger will be crossed "decades to centuries" ahead of when CO₂   increases (and pH decreases) alone would pose a threat to these bivalve larvae.

"At the current rate of change, there is not much more room for the waters off the Oregon coast to absorb more CO₂ without crossing the threshold we have identified with respect to saturation state," he said. Results of the study help explain commercial hatchery failures and why improving water chemistry in those hatcheries has been successful.

What kept the system more balanced in the geologic past likely included a combination of factors, the researchers say. One factor in past increases of carbon dioxide was high levels of volcanic activity. However, greater volcanic activity also coincides with more tectonic plate activity and uplift, increasing the weathering of rock surfaces - and thus alkalinity in rivers, where it eventually flowed into the ocean to offset the CO_2.

Computer models suggest that carbon dioxide is increasing through human activity some 100 to 1,000 times faster than the weathering processes that produce alkalinity can keep up, Waldbusser noted.

The Nature Climate Change study builds on previous research by Waldbusser and colleagues that outlined the mechanisms by which young bivalves create their shells after fertilization. In that study, the researchers found that young oysters and mussels had to build their shells within 48 hours to successfully begin feeding at a rate fast enough to survive, and that rate of shell-building would require a lot of energy. Thus in the presence of acidic water, they had to divert too much energy to shell-building and lacked the energy to swim and get food.

"The hatcheries call it the 'lazy larvae syndrome' because these tiny oysters just sink in the water and stop swimming," Waldbusser said. "These organisms have really sensitive windows to ocean acidification - even more sensitive then we previously thought."

In this latest study, the researchers used high-resolution images to analyze the development of oyster and mussel shells. They found that the organisms - which are about 1-100th the diameter of a human hair - actually build a complete calcium carbonate shell within six hours, about 12 hours after fertilization.

Throw off the ocean chemistry just a bit however, the researchers say, and a greater proportion of the shells do not develop normally. The ones that do are smaller, leading to potentially weaker organisms that will take longer to get to a size where they can settle into adult life.

"When the water is more saturated and has greater alkalinity it helps offset higher levels of carbon dioxide, ensuring that shell formation can proceed - and also making the shells bigger," Waldbusser said. "This can have a significant impact on their survivability into the future."

Shellfish hatcheries are altering their water chemistry based on the OSU research to create more favorable saturation state conditions for young bivalves; however this only helps organisms that can be cultured easily and increasing alkalinity in natural environments is a formidable challenge because of the amount required.

Other Oregon State researchers on the Nature Climate Change study included Burke Hales, Chris Langdon, Brian Haley, Paul Schrader, Elizabeth Brunner, Matthew Gray, Cale Miller and Iria Gimenez.