What the frog's eye tells the frog's brain
This is a key paper is biological systems theory. I took one of Jerome Letvin's courses at MIT, and later studied Ontological Design with Fernando Flores who pointed me to his teachers, Humberto Maturana and Francisco Varella. Maturana is a co-author on the paper.
Connecting back to semiotics, this cluster of thinkers and a network around them are probably the core of what is sometimes called "biologic semiotics or systems theory". I don't know the connection to Stafford Beer, but there has to be one if not only syncronicity.