FACT complex is involved in elongation and ensures fidelity in the initiation 
step of transcription by RNA polymerase (pol) II. Histone variant H2A.Z is found 
in nucleosomes at the 5'-end of many genes. We report here H2A.Z-chaperone 
activity of the yeast FACT complex on the short, nucleosome-free, non-coding, 
pol III-transcribed yeast tRNA genes. On a prototype gene, yeast SUP4, chromatin 
remodeler RSC and FACT regulate its transcription through novel mechanisms, 
wherein the two gene-flanking nucleosomes containing H2A.Z, play different 
roles. Nhp6, which ensures transcription fidelity and helps load yFACT onto the 
gene flanking nucleosomes, has inhibitory role. RSC maintains a nucleosome 
abutting the gene terminator downstream, which results in reduced transcription 
rate in active state while H2A.Z probably helps RSC in keeping the gene 
nucleosome-free and serves as stress-sensor. All these factors maintain an 
epigenetic state which allows the gene to return quickly from repressed to 
active state and tones down the expression from the active SUP4 gene, required 
probably to maintain the balance in cellular tRNA pool.