How does the net charge of proteins influence their elution order in anion-exchange chromatography?

Proteins with greater net positive charge elute first, while those with greater net negative charge elute last.

Proteins with greater net charge elute first, regardless of being positive or negative.

Proteins with neutral charge elute first, while those with any charge elute last.

Proteins with greater net negative charge elute first, while those with greater net positive charge elute last.