What do the SMTP error / reply codes mean?

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Here is a reference table of SMTP error / reply codes and a brief description of their meaning:

CodeDescription
211System status or system help reply.
214Help message.
220Domain service ready - ready to start TLS.
221Domain service closing transmission channel.
250OK - queuing for node started - requested mail action okay, completed.
251OK - no messages waiting for node - user not local will forward to forward path.
252OK - pending messages for node started - cannot VRFY user but will accept message and attempt delivery.
253OK - messages pending - messages for node started.
354Start mail input - end with ..
355Octet-offset is the transaction offset.
421Domain service not available - closing transmission channel.
432A password transition is needed.
450Requested mail action not taken: mailbox unavailable - ATRN request refused.
451Requested action aborted - local error in processing - unable to process ATRN request.
452Requested action not taken - insufficient system storage.
453You have no mail.
454TLS not available due to temporary reason - encryption is required for requested authentication mechanism.
458Unable to queue messages for node
459Node not allowed: reason.
500Command not recognized 'command' - syntax error.
501Syntax error - no parameters allowed.
502Command not implemented.
503Bad sequence of commands.
504Command parameter not implemented.
521Machine does not accept mail.
530Must issue a STARTTLS command first - encryption required for requested authentication mechanism.
534Authentication mechanism is too weak.
538Encryption required for requested authentication mechanism.
550Requested action not taken - mailbox unavailable.
551User not local - please try forward path.
552Requested mail action aborted - exceeded storage allocation.
553Requested action not taken - mailbox name not allowed.
554Transaction failed.

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