Chris, I went to the website you suggested. The "vacuolated nuclei" in the spermathecal lining of the drone-layer queens suggest apoptosis to me....meaning cell death. The nuclear appearance is classic for dead/dying cells. So, the spermathecal lining of drone-layer queens is unhealthy and may not be supporting the sperm very well. This might explain why those queens become drone layers....their sperm lose viability.
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