syntactic analysis - but learning that is dictated by their genetic programming, learning as thoroughly stereotyped as the most instinctive of behavioral responses

Ethologists are convinced that many animals survive through

learning—but learning that is dictated by their genetic programming,
learning as thoroughly stereotyped as the most instinctive of
behavioral responses.

This seems very convoluted structure to me. can anyone explain what is the relation between two parts of sentence before comma and after comma in the part "but learning that is dictated by their genetic programming, learning as thoroughly stereotyped as the most instinctive of behavioral responses". This part starting with the word "learning" before comma and after comma as well, what is their relation here?

Answer

"verb - but verb" is a common expression with scientific and technical writing (and also sometimes in literary criticism). It's a way for an author to create a more precise definition of a common word.

Everything that follows "- but learning" is restricting the normal definition of "learning" to the more precise, technical definition that the author wants to use. In this case, the author provides two parts to that definition ("dictated by genetic programming" and "thoroughly stereotyped..."). The comma you ask about just separates those two parts.

"learning - but learning ..." is a shorter way to say "learning: however, learning only in the narrow sense of ..."