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Your options are single click (thrust), and held click (pound)
Your upgrades and gameplan are tied to these interactions. Do not engage in either if a heart is about to pop (thrust has more "active frames") and DO be actively doing something with reversed hearts, when you unlock them.
Other than that, experiment and have fun!

(+2)

It still seems like the hearts take a significant penalty on you if you aren't doing anything when they explode. What's up with that?

(+1)

Hearts clearly seem to scale the further along you are in a run, certain monster hearts feeling absolutely egregious by loop 2 and the game as a whole being nigh impossible by four.
Sustain or big bonk is a necessity for every build because of it, and outside of some fine-tuning I sadly don't think there's much to be said about damage taken