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Photosynthetic complexes improve the transfer of excitation energy fromperipheral antennas to reaction centers in several ways. In particular, adownward energy funnel can direct excitons in the right direction, whilecoherent excitonic delocalization can enhance transfer rates through thecooperative phenomenon of supertransfer. However, isolating the role of purelycoherent effects is difficult because any change to the delocalization alsochanges the energy landscape. Here, we show that the relative importance of thetwo processes can be determined by comparing the natural light-harvestingapparatus with counterfactual models in which the delocalization and the energylandscape are altered. Applied to the example of purple bacteria, our approachshows that although supertransfer does enhance the rates somewhat, theenergetic funnelling plays the decisive role. Because delocalization has aminor role (and is sometimes detrimental), it is most likely not adaptive,being a side-effect of the dense chlorophyll packing that evolved to increaselight absorption per reaction center.

Distinguishing the roles of energy funnelling and delocalization in photosynthetic light harvesting