Hi Simon! Thanks for reading my essay and I have to say I'm a little starstruck because I'm a big fan of your work!
As to your question, I wanted to define meaning more along the lines of attention, I actually had Heidegger's notion of "care" in mind when I was trying to decide what made information meaninfgul. In biology, that's generically connected to survival but then, even for most animals it can be about much more then mere survival. Sexual selection, in particular, appears to be capable of creating very exotic forms of meaning, and for us most meaning is only dimly related to survival.
In essence my thinking was that mutual information was the first component of meaning, becuase a structure has to be capable of interacting with or detecting certain degrees of freedom in it's environment. Then I think the structure should also be capable of responding in a counterfactual manner, so if the environmental degrees of freedom being detected had taken a different value the structure would have computed a different response. That's how I would technically describe the act of paying attention to and responding to information, which is really what I wanted to say makes information meaningful. The e. coli example might have blurred that point since the bug is behaving that way "in order to" survive, but it's really the fact that it's sensing-computing-acting that causes the sensed degrees of freedom to be meaningful to the structure.
Finally then, as to the digital agent, if it ignores the symbol then the symbol is not meaningful to it, even if it dies as a result. If a large number of such agents are simulated and they also have some adaptive and reproductive capacity in their programing, then the ones that learn to pay attention to that symbol will in time take over the population. That's how I would depict the relationship between survival and meaning. Natural selection drives information increases in genomes at the population level by selecting for structures that have the fittest meaningful environmental relationships, meaning the ones that pay attention to what matters the best.
Hope that's clear and thanks again!
Joe