4.3.1: Official Definition
4.X.X
[RFC3463] (Standards Track) from the The Internet Engineering Task Force (IETF) definition for codes that start with 4.X.X:
A persistent transient failure is one in which the message as sent is valid, but persistence of some temporary condition has caused abandonment or delay of attempts to send the message. If this code accompanies a delivery failure report, sending in the future may be successful.
X.3.1
[RFC3463] (Standards Track) from the The Internet Engineering Task Force (IETF) definition for codes that end with X.3.1:
Mail system storage has been exceeded. The general semantics imply that the individual recipient may not be able to delete material to make room for additional messages. This is useful only as a persistent transient error.