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<</Limits[(springernature_natrevmolcellbio_769.indd:43.	Loffer, A. et al. A DCL3 dicing code within Pol IV-RDR2 transcripts div\
ersifies the siRNA pool guiding RNA-directed DNA methylation. eLife 11, e73260 \(2022\).\r:143)(springernature_natrevmolcellbio_769.indd:Fig. 4 | DNA methylation dynamics during plant reproduction. a, Tape\
tal cell-derived mobile small interfering RNAs \(siRNAs\) promote DNA methylation in neighbouring pollen mother cells \(PMCs\). In\
 tapetal cells, the production of 24-nucleot:444)]/Names[(springernature_natrevmolcellbio_769.indd:43.	Loffer, A. et al. A DCL3 dicing code within Pol IV-RDR2 transcripts div\
ersifies the siRNA pool guiding RNA-directed DNA methylation. eLife 11, e73260 \(2022\).\r:143)[2429 0 R/XYZ 39 577 null](springernature_natrevmolcellbio_769.indd:44.	Wang, Q. et al. Mechanism of siRNA production by a plant Dicer-RNA comp\
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late active DNA demethylation in Arabidopsis. Proc. Natl Acad. Sci. USA 116, 16641 16650 \(2019\).\r:243)[2429 0 R/XYZ 305 201 null](springernature_natrevmolcellbio_769.indd:Box 1 | Nomenclature and general conformation of plant nuclear RNA polymerase\
s\r \rDNA-dependent RNA polymerases Pol IV and Pol V evolved from Pol II293. All plant nuclear RNA polymerases contain 12\
 subunits. These subunits are named by the :118)[1521 0 R/XYZ 44 677 null](springernature_natrevmolcellbio_769.indd:Fig. 1 | Mechanism of RNA-directed DNA methylation. a, The canonical \
model of RNA-directed DNA methylation \(RdDM\). SHH1 preferentially binds to dimethylated histone 3 lysine 9 \(H3K9me2\) and promo\
tes DNA-dependent RNA polymerase IV \(Pol IV\):88)[300 0 R/XYZ 39 356 null](springernature_natrevmolcellbio_769.indd:Fig. 2 | Mechanism of active DNA demethylation. a, Recruitment of the\
 5-methylcytosine \(5mC\) DNA glycosylase ROS1 to its target loci requires DNA methylation readers. The IDM complex binds to meth\
ylated DNA \(red circles\) through its subuni:214)[2162 0 R/XYZ 305 680 null](springernature_natrevmolcellbio_769.indd:Fig. 3 | DNA methylation in transcription regulation. a, DNA methylat\
ion on genes and transposable elements \(TEs\) in the Arabidopsis thaliana genome mainly exists in three patterns. Fully silenc\
ed TEs and genes are associated with high l:289)[851 0 R/XYZ 39 136 null](springernature_natrevmolcellbio_769.indd:Fig. 4 | DNA methylation dynamics during plant reproduction. a, Tapet\
al cell-derived mobile small interfering RNAs \(siRNAs\) promote DNA methylation in neighbouring pollen mother cells \(PMCs\). In \
tapetal cells, the production of 24-nucleoti:407)[1777 0 R/XYZ 305 376 null](springernature_natrevmolcellbio_769.indd:Fig. 5 | Spontaneous epimutations and epigenome editing tools. a c\
,Spontaneous epimutations in toadflax \(Linaria vulgaris\), tomato and oil palm. a, The peloric mutant of toadflax has rad\
ially symmetrical flowers that are correlat:489)[1552 0 R/XYZ 39 195 null](springernature_natrevmolcellbio_769.indd:110.	Feng, Z. et al. Genetic analysis implicates a molecular chaperone com\
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e DNA demethylation to limit DNA hyper-methylation and transcriptional gene silencing. Mol. Cell 57, 971 983 \(2015\).\r:306)[2438 0 R/XYZ 39 625 null](springernature_natrevmolcellbio_769.indd:113.	Boone, B. A. et al. ACD15, ACD21, and SLN regulate the accumulation a\
nd mobility of MBD6 to silence genes and transposable elements. Sci. Adv. 9, eadi9036 \(2023\).\r:308)[2438 0 R/XYZ 39 585 null](springernature_natrevmolcellbio_769.indd:138.	Lang, Z. et al. Critical roles of DNA demethylation in the activation\
 of ripening-induced genes and inhibition of ripening-repressed genes in tomato fruit. Proc. Natl Acad. Sci. USA 114, E4511 \
E4519 \(2017\).\r:421)[2438 0 R/XYZ 39 137 null](springernature_natrevmolcellbio_769.indd:164.	Zhang, Y. Z. et al. Genome-wide distribution and functions of the AAE\
 complex in epigenetic regulation in Arabidopsis. J. Integr. Plant Biol. 63, 707 722 \(2021\).\r:364)[2438 0 R/XYZ 305 297 null](springernature_natrevmolcellbio_769.indd:168.	Walker, J. et al. Sexual-lineage-specific DNA methylation regulates m\
eiosis in Arabidopsis. Nat. Genet. 50, 130 137 \(2018\).\r:408)[2438 0 R/XYZ 305 233 null](springernature_natrevmolcellbio_769.indd:169.	Ong-Abdullah, M. et al. Loss of Karma transposon methylation underlie\
s the mantled somaclonal variant of oil palm. Nature 525, 533 537 \(2015\).\r:524)[2438 0 R/XYZ 305 217 null](springernature_natrevmolcellbio_769.indd:174.	She, W. et al. Chromatin reprogramming during the somatic-to-reproduc\
tive cell fate transition in plants. Development 140, 4008 4019 \(2013\).\r:389)[2438 0 R/XYZ 305 129 null](springernature_natrevmolcellbio_769.indd:191.	Long, J. et al. Nurse cell derived small RNAs define paternal epigene\
tic inheritance in Arabidopsis. Science 373, eabh0556 \(2021\).\r:412)[2456 0 R/XYZ 39 425 null](springernature_natrevmolcellbio_769.indd:213.	Zhou, S. et al. DNA demethylases remodel DNA methylation in rice game\
tes and zygote and are required for reproduction. Mol. Plant 14, 1569 1583 \(2021\).\r:456)[2456 0 R/XYZ 305 681 null](springernature_natrevmolcellbio_769.indd:30.	Huang, K. et al. Pol IV and RDR2: a two-RNA-polymerase machine that pr\
oduces double-stranded RNA. Science 374, 1579 1586 \(2021\).\r:123)[2384 0 R/XYZ 305 191 null](springernature_natrevmolcellbio_769.indd:43.	Loffer, A. et al. A DCL3 dicing code within Pol IV-RDR2 transcripts di\
versifies the siRNA pool guiding RNA-directed DNA methylation. eLife 11, e73260 \(2022\).\r:147)[2429 0 R/XYZ 39 577 null](springernature_natrevmolcellbio_769.indd:55.	Xie, G. et al. Structure and mechanism of the plant RNA polymerase V. 
Science 379, 1209 1213 \(2023\).\r:163)[2429 0 R/XYZ 39 353 null](springernature_natrevmolcellbio_769.indd:62.	Liu, W. et al. RNA-directed DNA methylation involves co-transcriptiona\
l small-RNA-guided \nslicing of polymerase V transcripts in Arabidopsis. Nat. Plants 4, 181 188 \(2018\).\r:180)[2429 0 R/XYZ 39 225 null](springernature_natrevmolcellbio_769.indd:63.	Wang, F., Huang, H. Y., Huang, J., Singh, J. & Pikaard, C. S. Enzymati\
c reactions of AGO4 in RNA-directed DNA methylation: siRNA duplex loading, passenger strand elimination, target RNA slicing, and\
 sliced target retention. Genes Dev. 37:182)[2429 0 R/XYZ 39 209 null](springernature_natrevmolcellbio_769.indd:67.	Zhang, H., Lang, Z. & Zhu, J. K. Dynamics and function of DNA methylat\
ion in plants. Nat. Rev. Mol. Cell Biol. 19, 489 506 \(2018\).\r:491)[2429 0 R/XYZ 39 137 null](springernature_natrevmolcellbio_769.indd:94.	Du, X. et al. Molecular basis of the plant ROS1-mediated active DNA de\
methylation. Nat. Plants 9, 271 279 \(2023\).\r:236)[2429 0 R/XYZ 305 289 null](springernature_natrevmolcellbio_769.indd:98.	Qian, W. et al. A histone acetyltransferase regulates active DNA demet\
hylation in Arabidopsis. Science 336, 1445 1448 \(2012\).\r:245)[2429 0 R/XYZ 305 217 null](springernature_natrevmolcellbio_769.indd:99.	Nie, W. F. et al. Histone acetylation recruits the SWR1 complex to reg\
ulate active DNA demethylation in Arabidopsis. Proc. Natl Acad. Sci. USA 116, 16641 16650 \(2019\).\r:247)[2429 0 R/XYZ 305 201 null](springernature_natrevmolcellbio_769.indd:Box 1 | Nomenclature and general conformation of plant nuclear RNA polymeras\
es\r \rDNA-dependent RNA polymerases Pol IV and Pol V evolved from Pol II293. All plant nuclear RNA polymerases contain 1\
2 subunits. These subunits are named by the:122)[1521 0 R/XYZ 44 677 null](springernature_natrevmolcellbio_769.indd:Fig. 1 | Mechanism of RNA-directed DNA methylation. a, The canonical\
 model of RNA-directed DNA methylation \(RdDM\). SHH1 preferentially binds to dimethylated histone 3 lysine 9 \(H3K9me2\) and prom\
otes DNA-dependent RNA polymerase IV \(Pol IV:94)[300 0 R/XYZ 39 356 null](springernature_natrevmolcellbio_769.indd:Fig. 2 | Mechanism of active DNA demethylation. a, Recruitment of th\
e 5-methylcytosine \(5mC\) DNA glycosylase ROS1 to its target loci requires DNA methylation readers. The IDM complex binds to met\
hylated DNA \(red circles\) through its subun:244)[2162 0 R/XYZ 305 680 null](springernature_natrevmolcellbio_769.indd:Fig. 3 | DNA methylation in transcription regulation. a, DNA methyla\
tion on genes and transposable elements \(TEs\) in the Arabidopsis thaliana genome mainly exists in three patterns. Fully silen\
ced TEs and genes are associated with high :296)[851 0 R/XYZ 39 136 null](springernature_natrevmolcellbio_769.indd:Fig. 4 | DNA methylation dynamics during plant reproduction. a, Tape\
tal cell-derived mobile small interfering RNAs \(siRNAs\) promote DNA methylation in neighbouring pollen mother cells \(PMCs\). In\
 tapetal cells, the production of 24-nucleot:444)[1777 0 R/XYZ 305 376 null]]>>
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,Spontaneous epimutations in toadflax \(Linaria vulgaris\), tomato and oil palm. a, The peloric mutant of toadflax has ra\
dially symmetrical flowers that are correla:494)(springernature_natrevmolcellbio_769.indd:Fig. 2 | Mechanism of active DNA demethylation. a, Recruitment o\
f the 5-methylcytosine \(5mC\) DNA glycosylase ROS1 to its target loci requires DNA methylation readers. The IDM complex binds to\
 methylated DNA \(red circles\) through its s:259)]/Names[(springernature_natrevmolcellbio_769.indd:Fig. 5 | Spontaneous epimutations and epigenome editing tools. a c
,Spontaneous epimutations in toadflax \(Linaria vulgaris\), tomato and oil palm. a, The peloric mutant of toadflax has ra\
dially symmetrical flowers that are correla:494)[1552 0 R/XYZ 39 195 null](springernature_natrevmolcellbio_769.indd:110.	Feng, Z. et al. Genetic analysis implicates a molecular chaperone co\
mplex in regulating epigenetic silencing of methylated genomic regions. J. Integr. Plant Biol. 63, 1451 1461 \(2021\).\r:303)[2438 0 R/XYZ 39 649 null](springernature_natrevmolcellbio_769.indd:111.	Lang, Z. et al. The methyl-CpG-binding protein MBD7 facilitates acti\
ve DNA demethylation to limit DNA hyper-methylation and transcriptional gene silencing. Mol. Cell 57, 971 983 \(2015\).\r:30\
9)[2438 0 R/XYZ 39 625 null](springernature_natrevmolcellbio_769.indd:113.	Boone, B. A. et al. ACD15, ACD21, and SLN regulate the accumulation \
and mobility of MBD6 to silence genes and transposable elements. Sci. Adv. 9, eadi9036 \(2023\).\r:310)[2438 0 R/XYZ 39 585 null](springernature_natrevmolcellbio_769.indd:138.	Lang, Z. et al. Critical roles of DNA demethylation in the activatio\
n of ripening-induced genes and inhibition of ripening-repressed genes in tomato fruit. Proc. Natl Acad. Sci. USA 114, E4511\
 E4519 \(2017\).\r:477)[2438 0 R/XYZ 39 137 null](springernature_natrevmolcellbio_769.indd:164.	Zhang, Y. Z. et al. Genome-wide distribution and functions of the AA\
E complex in epigenetic regulation in Arabidopsis. J. Integr. Plant Biol. 63, 707 722 \(2021\).\r:370)[2438 0 R/XYZ 305 297 null](springernature_natrevmolcellbio_769.indd:168.	Walker, J. et al. Sexual-lineage-specific DNA methylation regulates \
meiosis in Arabidopsis. Nat. Genet. 50, 130 137 \(2018\).\r:410)[2438 0 R/XYZ 305 233 null](springernature_natrevmolcellbio_769.indd:191.	Long, J. et al. Nurse cell derived small RNAs define paternal epigen\
etic inheritance in Arabidopsis. Science 373, eabh0556 \(2021\).\r:413)[2456 0 R/XYZ 39 425 null](springernature_natrevmolcellbio_769.indd:30.	Huang, K. et al. Pol IV and RDR2: a two-RNA-polymerase machine that p\
roduces double-stranded RNA. Science 374, 1579 1586 \(2021\).\r:125)[2384 0 R/XYZ 305 191 null](springernature_natrevmolcellbio_769.indd:55.	Xie, G. et al. Structure and mechanism of the plant RNA polymerase V.\
 Science 379, 1209 1213 \(2023\).\r:164)[2429 0 R/XYZ 39 353 null](springernature_natrevmolcellbio_769.indd:63.	Wang, F., Huang, H. Y., Huang, J., Singh, J. & Pikaard, C. S. Enzymat\
ic reactions of AGO4 in RNA-directed DNA methylation: siRNA duplex loading, passenger strand elimination, target RNA slicing, an\
d sliced target retention. Genes Dev. 3:183)[2429 0 R/XYZ 39 209 null](springernature_natrevmolcellbio_769.indd:94.	Du, X. et al. Molecular basis of the plant ROS1-mediated active DNA d\
emethylation. Nat. Plants 9, 271 279 \(2023\).\r:237)[2429 0 R/XYZ 305 289 null](springernature_natrevmolcellbio_769.indd:98.	Qian, W. et al. A histone acetyltransferase regulates active DNA deme\
thylation in Arabidopsis. Science 336, 1445 1448 \(2012\).\r:271)[2429 0 R/XYZ 305 217 null](springernature_natrevmolcellbio_769.indd:99.	Nie, W. F. et al. Histone acetylation recruits the SWR1 complex to re\
gulate active DNA demethylation in Arabidopsis. Proc. Natl Acad. Sci. USA 116, 16641 16650 \(2019\).\r:249)[2429 0 R/XYZ 305 201 null](springernature_natrevmolcellbio_769.indd:Fig. 1 | Mechanism of RNA-directed DNA methylation. a, The canonica\
l model of RNA-directed DNA methylation \(RdDM\). SHH1 preferentially binds to dimethylated histone 3 lysine 9 \(H3K9me2\) and pro\
motes DNA-dependent RNA polymerase IV \(Pol I:129)[300 0 R/XYZ 39 356 null](springernature_natrevmolcellbio_769.indd:Fig. 2 | Mechanism of active DNA demethylation. a, Recruitment of t\
he 5-methylcytosine \(5mC\) DNA glycosylase ROS1 to its target loci requires DNA methylation readers. The IDM complex binds to me\
thylated DNA \(red circles\) through its subu:246)[2162 0 R/XYZ 305 680 null](springernature_natrevmolcellbio_769.indd:Fig. 3 | DNA methylation in transcription regulation. a, DNA methyl\
ation on genes and transposable elements \(TEs\) in the Arabidopsis thaliana genome mainly exists in three patterns. Fully sile\
nced TEs and genes are associated with high:333)[851 0 R/XYZ 39 136 null](springernature_natrevmolcellbio_769.indd:Fig. 4 | DNA methylation dynamics during plant reproduction. a, Tap\
etal cell-derived mobile small interfering RNAs \(siRNAs\) promote DNA methylation in neighbouring pollen mother cells \(PMCs\). I\
n tapetal cells, the production of 24-nucleo:465)[1777 0 R/XYZ 305 376 null](springernature_natrevmolcellbio_769.indd:Fig. 5 | Spontaneous epimutations and epigenome editing tools. a \
c,Spontaneous epimutations in toadflax \(Linaria vulgaris\), tomato and oil palm. a, The peloric mutant of toadflax has r\
adially symmetrical flowers that are correl:525)[1552 0 R/XYZ 39 195 null](springernature_natrevmolcellbio_769.indd:110.	Feng, Z. et al. Genetic analysis implicates a molecular chaperone c\
omplex in regulating epigenetic silencing of methylated genomic regions. J. Integr. Plant Biol. 63, 1451 1461 \(2021\).\r:30\
7)[2438 0 R/XYZ 39 649 null](springernature_natrevmolcellbio_769.indd:111.	Lang, Z. et al. The methyl-CpG-binding protein MBD7 facilitates act\
ive DNA demethylation to limit DNA hyper-methylation and transcriptional gene silencing. Mol. Cell 57, 971 983 \(2015\).\r:3\
11)[2438 0 R/XYZ 39 625 null](springernature_natrevmolcellbio_769.indd:138.	Lang, Z. et al. Critical roles of DNA demethylation in the activati\
on of ripening-induced genes and inhibition of ripening-repressed genes in tomato fruit. Proc. Natl Acad. Sci. USA 114, E451\
1 E4519 \(2017\).\r:480)[2438 0 R/XYZ 39 137 null](springernature_natrevmolcellbio_769.indd:30.	Huang, K. et al. Pol IV and RDR2: a two-RNA-polymerase machine that \
produces double-stranded RNA. Science 374, 1579 1586 \(2021\).\r:127)[2384 0 R/XYZ 305 191 null](springernature_natrevmolcellbio_769.indd:55.	Xie, G. et al. Structure and mechanism of the plant RNA polymerase V\
. Science 379, 1209 1213 \(2023\).\r:166)[2429 0 R/XYZ 39 353 null](springernature_natrevmolcellbio_769.indd:94.	Du, X. et al. Molecular basis of the plant ROS1-mediated active DNA \
demethylation. Nat. Plants 9, 271 279 \(2023\).\r:238)[2429 0 R/XYZ 305 289 null](springernature_natrevmolcellbio_769.indd:98.	Qian, W. et al. A histone acetyltransferase regulates active DNA dem\
ethylation in Arabidopsis. Science 336, 1445 1448 \(2012\).\r:277)[2429 0 R/XYZ 305 217 null](springernature_natrevmolcellbio_769.indd:Fig. 1 | Mechanism of RNA-directed DNA methylation. a, The canonic\
al model of RNA-directed DNA methylation \(RdDM\). SHH1 preferentially binds to dimethylated histone 3 lysine 9 \(H3K9me2\) and pr\
omotes DNA-dependent RNA polymerase IV \(Pol :152)[300 0 R/XYZ 39 356 null](springernature_natrevmolcellbio_769.indd:Fig. 2 | Mechanism of active DNA demethylation. a, Recruitment of \
the 5-methylcytosine \(5mC\) DNA glycosylase ROS1 to its target loci requires DNA methylation readers. The IDM complex binds to m\
ethylated DNA \(red circles\) through its sub:248)[2162 0 R/XYZ 305 680 null](springernature_natrevmolcellbio_769.indd:Fig. 3 | DNA methylation in transcription regulation. a, DNA methy\
lation on genes and transposable elements \(TEs\) in the Arabidopsis thaliana genome mainly exists in three patterns. Fully sil\
enced TEs and genes are associated with hig:353)[851 0 R/XYZ 39 136 null](springernature_natrevmolcellbio_769.indd:Fig. 4 | DNA methylation dynamics during plant reproduction. a, Ta\
petal cell-derived mobile small interfering RNAs \(siRNAs\) promote DNA methylation in neighbouring pollen mother cells \(PMCs\). \
In tapetal cells, the production of 24-nucle:476)[1777 0 R/XYZ 305 376 null](springernature_natrevmolcellbio_769.indd:Fig. 5 | Spontaneous epimutations and epigenome editing tools. a 
c,Spontaneous epimutations in toadflax \(Linaria vulgaris\), tomato and oil palm. a, The peloric mutant of toadflax has \
radially symmetrical flowers that are corre:547)[1552 0 R/XYZ 39 195 null](springernature_natrevmolcellbio_769.indd:110.	Feng, Z. et al. Genetic analysis implicates a molecular chaperone \
complex in regulating epigenetic silencing of methylated genomic regions. J. Integr. Plant Biol. 63, 1451 1461 \(2021\).\r:3\
13)[2438 0 R/XYZ 39 649 null](springernature_natrevmolcellbio_769.indd:138.	Lang, Z. et al. Critical roles of DNA demethylation in the activat\
ion of ripening-induced genes and inhibition of ripening-repressed genes in tomato fruit. Proc. Natl Acad. Sci. USA 114, E45\
11 E4519 \(2017\).\r:482)[2438 0 R/XYZ 39 137 null](springernature_natrevmolcellbio_769.indd:30.	Huang, K. et al. Pol IV and RDR2: a two-RNA-polymerase machine that\
 produces double-stranded RNA. Science 374, 1579 1586 \(2021\).\r:132)[2384 0 R/XYZ 305 191 null](springernature_natrevmolcellbio_769.indd:94.	Du, X. et al. Molecular basis of the plant ROS1-mediated active DNA\
 demethylation. Nat. Plants 9, 271 279 \(2023\).\r:240)[2429 0 R/XYZ 305 289 null](springernature_natrevmolcellbio_769.indd:Fig. 1 | Mechanism of RNA-directed DNA methylation. a, The canoni\
cal model of RNA-directed DNA methylation \(RdDM\). SHH1 preferentially binds to dimethylated histone 3 lysine 9 \(H3K9me2\) and p\
romotes DNA-dependent RNA polymerase IV \(Pol:170)[300 0 R/XYZ 39 356 null](springernature_natrevmolcellbio_769.indd:Fig. 2 | Mechanism of active DNA demethylation. a, Recruitment of\
 the 5-methylcytosine \(5mC\) DNA glycosylase ROS1 to its target loci requires DNA methylation readers. The IDM complex binds to \
methylated DNA \(red circles\) through its su:251)[2162 0 R/XYZ 305 680 null](springernature_natrevmolcellbio_769.indd:30.	Huang, K. et al. Pol IV and RDR2: a two-RNA-polymerase machine tha\
t produces double-stranded RNA. Science 374, 1579 1586 \(2021\).\r:133)[2384 0 R/XYZ 305 191 null](springernature_natrevmolcellbio_769.indd:Fig. 2 | Mechanism of active DNA demethylation. a, Recruitment o\
f the 5-methylcytosine \(5mC\) DNA glycosylase ROS1 to its target loci requires DNA methylation readers. The IDM complex binds to\
 methylated DNA \(red circles\) through its s:259)[2162 0 R/XYZ 305 680 null]]>>
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