We envision multiple steps in telomere maintenance, based largely on geneti
c data from budding yeast. First, the telomere must unfold or open itself s
uch that the free end is accessible to the appropriate enzymatic machinery.
Second, telomerase must be recruited, together with the DNA replication ma
chinery that synthesizes the C-rich strand. The processivity of telomerase
is regulated both by a length-sensing feedback mechanism and by second-stra
nd synthesis. Finally, the telosome refolds into a protective end structure
. If telomerase is nonfunctional, recombination may occur once telomeres ar
e open. Multiple pathways regulate these different steps, producing a highl
y dynamic chromosomal cap.