Uneven crumb -- proofing or other problem?

I made these loaves based on the San Joaquin recipe by dmsnyder, but I upped the whole grain content to 50% (mostly wheat, but also a significant amount of rye), and raised the hydration (by an unmeasured amount, but it ended up feeling higher than I had intended). I only cut one of the loaves so far, but as you can see, the crumb was pretty irregular. Is this a sign of a particular part of the process I can improve on? Proofing? Something else? Or is it just normal variation? It seems a little too drastic for that last option.
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