The C1-FFL with an AND gate was shown to have a function of a ‘sign-sensitive delay’ element and a persistence detector both theoretically <ref name="man1"/> and experimentally<ref name="man2">{{cite journal |doi=10.1016/j.jmb.2003.09.049 |vauthors=Mangan S, Zaslaver A, Alon U |title=The coherent feedforward loop serves as a sign-sensitive delay element in transcription networks |journal=J. Mol. Biol. |volume=334 |issue=2 |pages=197–204 |date=November 2003 |pmid=14607112 |citeseerx=10.1.1.110.4629 }}</ref> with the arabinose system of ''[[Escherichia coli|E. coli]]''. This means that this motif can provide pulse filtration in which short pulses of signal will not generate a response but persistent signals will generate a response after short delay. The shut off of the output when a persistent pulse is ended will be fast. The opposite behavior emerges in the case of a sum gate with fast response and delayed shut off as was demonstrated in the flagella system of ''[[Escherichia coli|E. coli]]''.<ref name="kal1">{{cite journal |vauthors=Kalir S, Mangan S, Alon U |title=A coherent feed-forward loop with a SUM input function prolongs flagella expression in ''Escherichia coli'' |journal=Mol. Syst. Biol. |volume=1 |pages=E1–E6 |year=2005 |pmid=16729041 |pmc=1681456 |doi=10.1038/msb4100010 |issue=1}}</ref> De novo evolution of C1-FFLs in [[gene regulatory network]]s has been demonstrated computationally in response to selection to filter out an idealized short signal pulse, but for non-idealized noise, a dynamics-based system of feed-forward regulation with different topology was instead favored.<ref>{{cite journal |last1=Xiong |first1=Kun |last2=Lancaster |first2=Alex K. |last3=Siegal |first3=Mark L. |last4=Masel |first4=Joanna |title=Feed-forward regulation adaptively evolves via dynamics rather than topology when there is intrinsic noise |journal=Nature Communications |date=3 June 2019 |volume=10 |issue=1 |pages=2418 |doi=10.1038/s41467-019-10388-6|pmid=31160574 |pmc=6546794 }}</ref> | The C1-FFL with an AND gate was shown to have a function of a ‘sign-sensitive delay’ element and a persistence detector both theoretically <ref name="man1"/> and experimentally<ref name="man2">{{cite journal |doi=10.1016/j.jmb.2003.09.049 |vauthors=Mangan S, Zaslaver A, Alon U |title=The coherent feedforward loop serves as a sign-sensitive delay element in transcription networks |journal=J. Mol. Biol. |volume=334 |issue=2 |pages=197–204 |date=November 2003 |pmid=14607112 |citeseerx=10.1.1.110.4629 }}</ref> with the arabinose system of ''[[Escherichia coli|E. coli]]''. This means that this motif can provide pulse filtration in which short pulses of signal will not generate a response but persistent signals will generate a response after short delay. The shut off of the output when a persistent pulse is ended will be fast. The opposite behavior emerges in the case of a sum gate with fast response and delayed shut off as was demonstrated in the flagella system of ''[[Escherichia coli|E. coli]]''.<ref name="kal1">{{cite journal |vauthors=Kalir S, Mangan S, Alon U |title=A coherent feed-forward loop with a SUM input function prolongs flagella expression in ''Escherichia coli'' |journal=Mol. Syst. Biol. |volume=1 |pages=E1–E6 |year=2005 |pmid=16729041 |pmc=1681456 |doi=10.1038/msb4100010 |issue=1}}</ref> De novo evolution of C1-FFLs in [[gene regulatory network]]s has been demonstrated computationally in response to selection to filter out an idealized short signal pulse, but for non-idealized noise, a dynamics-based system of feed-forward regulation with different topology was instead favored.<ref>{{cite journal |last1=Xiong |first1=Kun |last2=Lancaster |first2=Alex K. |last3=Siegal |first3=Mark L. |last4=Masel |first4=Joanna |title=Feed-forward regulation adaptively evolves via dynamics rather than topology when there is intrinsic noise |journal=Nature Communications |date=3 June 2019 |volume=10 |issue=1 |pages=2418 |doi=10.1038/s41467-019-10388-6|pmid=31160574 |pmc=6546794 }}</ref> |