Alex worked extensively on the flash-induced ECS that indicates the delocalized trans-thylakoid electric potential difference, his first paper dealing with electrogenic events (fast and slow) in chloroplasts and their relation to the ECS (Hope and Morland 1980). In particular, he used the “slow” rise of the ECS, together with the concomitant reduction of cyt b to establish, with others, the working of a “Q-cycle” as originally

proposed for mitochondria by Peter Mitchell, under most conditions (Hope 1993). He confirmed the Q-cycle as normally occurring in isolated thylakoids (Hope 1993) and in intact leaves (Chow and Hope 2004b). The “fast” rise of the ECS indicates delocalized charge separation across the thylakoid membrane at the two photosystems.

By progressively photoinactivating PS II and extrapolating to zero functional PS II, Alex proposed, one could obtain the BV-6 separate contribution from PS I, and hence determine the often controversial PS II:PS I stoichiometry (Chow and Hope 1998; Fan et al. 2007a). Similarly, when charge separation in PS I is hampered by the photo-oxidation of P700 in the reaction centre by steady background far-red light, the separate contribution of PS II could be obtained after accounting for a small level of reduced P700 remaining. This approach, too, could be used to determine the photosystem stoichiometry in leaves (Fan BI 10773 et al. 2007a). Alex continually attempted to design equipment that had superior signal-to-noise ratio. His extensive measurements of the kinetics of electron transfers around the cyt bf complex using both isolated chloroplasts and isolated macromolecular complexes from thylakoids, Galactosylceramidase laid the groundwork for a full mathematical description of these processes (Hope 2000). With collaborators, he made use of the Inverse Method to optimize estimation of kinetic Belnacasan cell line parameters in electron transfers around the cyt bf complex (Hope et al. 1992). He set up a minimal set of reactions with differential equations to describe the rates of variation of the concentration

of all relevant species in terms of the rate coefficients of the reactions. He used the Inverse Method as a means of objectively optimizing the rate coefficients by systematically varying them while comparing model data with the corresponding experimental data until some specified minimum error integral was reached. The result of this process was to arrive at some new rate coefficients for thylakoids, with varying degrees of precision. He further examined the kinetic constants for the electron transfer reactions in thylakoids between plastocyanin and cyt f in cyt bf complexes, and between plastocyanin and the reaction centre of PS I, altering the parameters through changes in pH or ionic strength (Hope 2000) or hydrostatic pressure.