Elevated pH and Availability of Carbon Dioxide
The first possible mechanism whereby dinoflagellates may respond negatively to pH relates to their acquisition of carbon for photosynthesis. All photosynthesizing organisms need to take up carbon dioxide in some fashion in order to use it to make organic molecules. In a previous article I detailed many of these mechanisms, and they include a variety of different ways of taking carbon dioxide or bicarbonate/carbonate from the water and into the organism.
As the pH is raised at constant carbonate alkalinity, the amount of carbon dioxide in the water declines. A rise in pH of 0.3 units implies approximately a 50% reduction in the available carbon dioxide, but not a significant decrease in bicarbonate (or carbonate). Some organisms are known to suffer considerably from this loss in available carbon dioxide, particularly those that do not use bicarbonate or carbonate. Some species of macroalgae, for example, can photosynthesize only 18% as fast at pH 8.7 as they do at pH 8.1, while others do just as well at the higher pH.
So the question here is whether the problem dinoflagellates have this same response or not. As mentioned above, the exact species that are a problem in reef aquaria have not been identified, and even if identified, have probably not been studied with respect to their pH response. From the literature, some dinoflagellates can take up carbon dioxide only as carbon dioxide, while others can use bicarbonate.
Two marine dinoflagellates, Amphidinium carterae Hulburt and Heterocapsa oceanica Stein, demonstrate active uptake of carbon dioxide (or carbonic acid), but not bicarbonate. Because this mechanism is fundamentally limited in its effectiveness, it has been speculated that these organisms may be CO2-limited in their natural environment.5 These species would likely be stressed considerably if the pH of a reef aquarium containing them were raised substantially. On the other hand, three marine bloom-forming (red tide) dinoflagellates, Prorocentrum minimum, Heterocapsa triquetra and Ceratium lineatum,6 have been shown to take up bicarbonate directly, with bicarbonate accounting for approximately 80% of the carbon dioxide they use in photosynthesis. It is believed that these dinoflagellates are not carbon limited in photosynthesis due to their efficient direct bicarbonate uptake mechanisms, so they may not be overly stressed (by this mechanism) by raising the pH to levels achievable in a reef aquarium.
Dinoflagellates' Internal pH
Organisms typically have strong control of their internal pH regardless of small changes in the external pH. Internal cellular pH is often near pH 7. The green alga Chlorella saccharophila, for example, has an internal pH of 7.3 that does not change across the external pH range from pH 5 to pH 7.5. As the pH drops below 5, however, its internal pH begins to drop and falls to 6.4 when the external pH reaches 3.0.7
The reason that organisms control their internal pH so strongly is that the rate of many different biochemical processes depends on pH. Enzymes, for example, catalyze reactions, and their ability to do so nearly always depends on the pH. So, in order to ensure that the myriad chemical reactions taking place inside cells operate at desirable rates, organisms keep their internal pH from fluctuating. If their internal pH strayed significantly from "normal" for that organism, chemical imbalances are likely to arise and the organism can be significantly stressed.
As an aside, the primary reason that I believe that small sudden pH changes do not stress most reef aquarium organisms, as long as the pH does not move outside the normal pH range that is acceptable to them, is because of this strong internal pH control. For example, I do not believe that a sudden rise in pH from 8.1 to 8.4 is any more stressful for most marine organisms than is a stable continuous pH 8.1 or 8.4.