My assessment is based souly off the information provided.
Swords are the only plant I noticed in the picture, and they do well without CO2 supplementation, and also do well in lower to moderate light. Plus they're heavy root feeders - a very small amount of nutrients is absorbed through the water column(in this case) - gravel is not good for holding nutrients, and requires root tabs to help combat that at a minimum.
However, your post is on point and accurate. Could be any number of things, and more information would be required to give a more percise answer. :-)






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