We present a model of the S-state mechanism, as well as an improved ei
genvalue analysis, that integrate into a coherent ensemble several fea
tures found since the S-state model was initially developed. These fea
tures include the presence of S-1, deactivations in the dark interval
between flashes, and the change in the number of active PS II centers
by photoinhibition or photoactivation. A new feature is the capacity t
o predict the steady-state distribution of S-states under conditions o
f steady photoinhibition or photoactivation. The improved eigenvalue a
nalysis allowed the calculation of the initial S-state distribution. I
n addition, the model resolved 'true' photochemical misses from appare
nt misses due to deactivations in the dark interval between flashes. T
he model suggested that most of the misses that are commonly reported
are due to deactivations, and not to an intrinsic inefficiency of the
photochemical mechanism of PS II. Because models that allow double-hit
s encompassing the S-2 to S-3 transition Often predict negative initia
l quantities of S-2 in cyanobacteria, our proposed model specifically
prohibited them. The model accounts for inhomogeneous misses and a ste
ady-state distribution of the type (S-2)approximate to(S-1)>(S-3)appro
ximate to(S-0). This 5-step model uses only 4 probabilities, and is th
erefore easy to handle. The use of this model is critical for the anal
ysis of several cyanobacterial strains, as well as for any species tha
t show non-negligible deactivations in the dark interval between flash
es.