HEN1

UniProt ID: Q9C5Q8
Organism: Arabidopsis thaliana
Review Status: COMPLETE
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Gene Description

HEN1 is the Arabidopsis SAM-dependent small RNA 3' terminal 2'-O-methyltransferase. It methylates the 2'-OH of the 3'-terminal ribose on miRNA/miRNA-star and siRNA/siRNA-star duplexes after Dicer-like processing, protecting small RNAs from uridylation, trimming, and degradation. HEN1 acts in the DCL1/HYL1-associated small-RNA maturation context and supports miRNA, siRNA, ta-siRNA, and antiviral silencing outputs; floral and leaf patterning phenotypes are downstream consequences of destabilized small RNAs.

Proposed New Ontology Terms

small RNA 3-prime-end protection by 2-prime-O-methylation

Definition: A small-RNA metabolic process in which methylation of the 2-prime-OH of the 3-prime-terminal ribose protects mature small RNAs from uridylation, trimming, and exonucleolytic decay.

Justification: HEN1's direct molecular function is captured by small RNA 2-prime-O-ribose methyltransferase activity, but the current core-function summary also needs to distinguish the downstream protective effect of that modification. A process term for small-RNA 3-prime-end protection would avoid reusing the same catalytic molecular-function term for both methyl transfer and stabilization.

Parent term: RNA processing

Existing Annotations Review

GO Term Evidence Action Reason
GO:0034587 piRNA processing
IBA
GO_REF:0000033
REMOVE
Summary: HEN1-family proteins include animal piRNA methyltransferases, but Arabidopsis HEN1 methylates plant miRNA and siRNA duplexes.
Reason: piRNA processing is an over-transfer from metazoan HEN1/HENMT1 family context and is not supported for Arabidopsis HEN1.
Propagation Review
Root cause: PROPAGATION BAD
Failure modes: LINEAGE OR TAXON MISMATCH
Sources checked:
PANTHER:PTN000483941 Β· HEN1/HENMT1 family node SUPPORTS SOURCE BUT NOT TARGET
The piRNA-processing IBD is seeded exclusively by metazoan HEN1/HENMT1 orthologs (fly, mouse, worm, zebrafish) but placed at a Eukaryota-wide node; the piRNA pathway does not exist in plants, so the term cannot apply to Arabidopsis HEN1. A taxon constraint or shallower IBD placement would fix this at source.
MGI:MGI:1913965 Β· mouse Henmt1 SUPPORTS SOURCE BUT NOT TARGET
Genuine piRNA 2-prime-O-methyltransferase seed; the piRNA context is metazoan-specific.
GO:0008171 O-methyltransferase activity
IBA
GO_REF:0000033
MODIFY
Summary: HEN1 is an RNA methyltransferase, but O-methyltransferase activity is less specific than the small-RNA 2-prime-O-ribose methyltransferase term.
Reason: Use the small-RNA terminal 2-prime-O-methyltransferase term to capture HEN1 substrate and reaction specificity.
Propagation Review
Root cause: TERM SCOPING PROBLEM
Failure modes: GRANULARITY MISMATCH
Sources checked:
PANTHER:PTN000483941 Β· HEN1/HENMT1 family node SUPPORTS TRANSFER
All seeds are small-RNA terminal 2-prime-O-methyltransferases; the generic O-methyltransferase term under-specifies the shared activity, hence the refinement.
GO:0003755 peptidyl-prolyl cis-trans isomerase activity
IEA
GO_REF:0000002
REMOVE
Summary: HEN1 includes a PPIase-like/FKBP-like domain, but its curated function is small-RNA methyltransferase activity.
Reason: The PPIase-like/FKBP-like domain is present, but no evidence supports HEN1 peptidyl-prolyl isomerase catalytic activity; the supported biochemical function is small-RNA methyltransferase activity.
GO:0005634 nucleus
IEA
GO_REF:0000044
ACCEPT
Summary: HEN1 is associated with the nuclear DCL1/HYL1 small-RNA maturation context.
Reason: The best-supported site for HEN1 action is nuclear small-RNA processing/methylation before or around AGO loading.
Supporting Evidence:
file:ARATH/HEN1/HEN1-deep-research-falcon.md
Direct interaction mapping and mechanistic models place HEN1 in the nuclear miRNA-processing context.
GO:0008171 O-methyltransferase activity
IEA
GO_REF:0000002
MODIFY
Summary: HEN1 is an RNA methyltransferase, but O-methyltransferase activity is less specific than the small-RNA 2-prime-O-ribose methyltransferase term.
Reason: Use the small-RNA terminal 2-prime-O-methyltransferase term to capture HEN1 substrate and reaction specificity.
GO:0008173 RNA methyltransferase activity
IEA
GO_REF:0000002
MODIFY
Summary: HEN1 is an RNA methyltransferase, but RNA methyltransferase activity is less specific than the small-RNA 2-prime-O-ribose methyltransferase term.
Reason: Use the small-RNA terminal 2-prime-O-methyltransferase term to capture HEN1 substrate and reaction specificity.
GO:0016070 RNA metabolic process
IEA
GO_REF:0000117
MODIFY
Summary: HEN1 participates in small-RNA metabolism by methylating mature miRNA/siRNA duplexes.
Reason: RNA metabolic process is too broad; small-RNA methylation and miRNA/siRNA processing-stabilization outputs are more informative.
GO:0090486 small RNA 2'-O-ribose methyltransferase activity
IEA
GO_REF:0000003
ACCEPT
Summary: HEN1 catalyzes terminal 2-prime-O-ribose methylation of plant small RNAs. The HEN1 crystal structure (PDB 3HTX) shows the SAH cofactor product bound in the Rossmann MTase domain as in classical SAM-dependent methyltransferases, with a catalytic Mg ion coordinating the 3-prime-terminal 2-prime and 3-prime hydroxyls, consistent with 2-prime-O-ribose methyl transfer.
Reason: This is the most specific molecular-function term for Arabidopsis HEN1.
Supporting Evidence:
file:ARATH/HEN1/HEN1-deep-research-falcon.md
HEN1 is a small RNA 3-prime-terminal 2-prime-O-methyltransferase.
file:ARATH/HEN1/HEN1-uniprot.txt
Reaction=small RNA 3'-end nucleotide + S-adenosyl-L-methionine = small RNA 3'-end 2'-O-methylnucleotide + S-adenosyl-L-homocysteine + H(+);
file:interpro/panther/PTHR21404/PTHR21404-entries.csv
Q9C5Q8,Small RNA 2'-O-methyltransferase
PMID:19812675
The MTase domain of HEN1 adopts a core Ξ±/Ξ² Rossmann structure, in which the cofactor product AdoHcy is bound as in classical S-adenosyl-l-methionine (AdoMet)-dependent MTases
PMID:19812675
both the 2β€² and 3β€² hydroxyls of G22m and the side chains of four invariant residues (E796, E799, H800 and H860) are coordinated to a metal ion, Mg2+
GO:0090486 small RNA 2'-O-ribose methyltransferase activity
IEA
GO_REF:0000116
ACCEPT
Summary: HEN1 catalyzes terminal 2-prime-O-ribose methylation of plant small RNAs. The 3.1 A crystal structure of full-length Arabidopsis HEN1 (PDB 3HTX) bound to a 22-nucleotide small RNA duplex with the cofactor product SAH (S-adenosyl-L-homocysteine) and Mg directly visualizes the substrate as a small-RNA duplex and confirms terminal 2-prime-O-methyltransferase chemistry.
Reason: This is the most specific molecular-function term for Arabidopsis HEN1.
Supporting Evidence:
file:ARATH/HEN1/HEN1-deep-research-falcon.md
HEN1 is a small RNA 3-prime-terminal 2-prime-O-methyltransferase.
file:ARATH/HEN1/HEN1-uniprot.txt
Reaction=small RNA 3'-end nucleotide + S-adenosyl-L-methionine = small RNA 3'-end 2'-O-methylnucleotide + S-adenosyl-L-homocysteine + H(+);
file:interpro/panther/PTHR21404/PTHR21404-entries.csv
Q9C5Q8,Small RNA 2'-O-methyltransferase
PMID:19812675
report the 3.1 A crystal structure of full-length HEN1 from Arabidopsis in complex with a 22-nucleotide small RNA duplex and cofactor product S-adenosyl-l-homocysteine
PMID:19812675
A subset of small RNAs, such as microRNAs and small interfering RNAs (siRNAs) in plants, Piwi-interacting RNAs in animals and siRNAs in Drosophila, requires an additional crucial step for their maturation; that is, 2'-O-methylation on the 3' terminal nucleotide. A conserved S-adenosyl-l-methionine-dependent RNA methyltransferase, HUA ENHANCER 1 (HEN1), and its homologues are responsible for this specific modification.
GO:0010267 ta-siRNA processing
IMP
PMID:15469823
Endogenous trans-acting siRNAs regulate the accumulation of ...
KEEP AS NON CORE
Summary: HEN1 supports ta-siRNA processing by methylating and stabilizing the small-RNA duplexes that feed these pathways.
Reason: HEN1 is required for stable miRNA/siRNA pathway outputs, but the direct catalytic step is small-RNA 2-prime-O-methylation rather than Dicer cleavage or AGO-mediated target repression.
Supporting Evidence:
file:ARATH/HEN1/HEN1-uniprot.txt
Can methylate 3'-end of microRNAs (miRNAs), small interfering RNAs (siRNas) and trans-acting small interfering RNAs (ta-siRNAs).
GO:0005737 cytoplasm
ISM
GO_REF:0000122
UNDECIDED
Summary: The AtSubP-inferred cytoplasm row lacks positive experimental support for HEN1 cytoplasmic localization.
Reason: HEN1's best-supported context is small-RNA methylation around nuclear miRNA/siRNA maturation; downstream cytoplasmic small-RNA turnover is not sufficient evidence for a cytoplasm localization annotation.
GO:0010305 leaf vascular tissue pattern formation
IMP
PMID:22623415
The microRNA pathway genes AGO1, HEN1 and HYL1 participate i...
KEEP AS NON CORE
Summary: hen1 mutants affect leaf vascular tissue pattern formation.
Reason: The developmental process is a downstream phenotype of altered miRNA/siRNA stability rather than a direct HEN1 function.
GO:0010589 leaf proximal/distal pattern formation
IMP
PMID:22623415
The microRNA pathway genes AGO1, HEN1 and HYL1 participate i...
KEEP AS NON CORE
Summary: hen1 mutants affect leaf proximal/distal pattern formation.
Reason: The developmental process is a downstream phenotype of altered miRNA/siRNA stability rather than a direct HEN1 function.
GO:0009616 RNAi-mediated antiviral immune response
IMP
PMID:17090584
Four plant Dicers mediate viral small RNA biogenesis and DNA...
KEEP AS NON CORE
Summary: HEN1 contributes to virus-induced silencing and viral-small-RNA stability through terminal small-RNA methylation.
Reason: The antiviral role is a pathway output of HEN1-mediated small-RNA stabilization; the direct catalytic function remains small-RNA 2-prime-O-methylation rather than Dicing or Argonaute effector activity.
Supporting Evidence:
PMID:17090584
DCL4 in conjunction with RDR6 and HEN1 specifically facilitates extensive virus-induced silencing in new growth.
GO:0005634 nucleus
IDA
PMID:17442570
Identification of nuclear dicing bodies containing proteins ...
ACCEPT
Summary: The cited D-body paper places plant miRNA processing in the nucleus but directly images DCL1/HYL1/SE rather than HEN1.
Reason: Although PMID:17442570 directly images other D-body proteins, HEN1 has independent support for the same nuclear miRNA-processing context, so the nucleus annotation is retained consistently.
Supporting Evidence:
file:ARATH/HEN1/HEN1-deep-research-falcon.md
Direct interaction mapping and mechanistic models place HEN1 in the nuclear miRNA-processing context.
GO:0005737 cytoplasm
IDA
PMID:17442570
Identification of nuclear dicing bodies containing proteins ...
UNDECIDED
Summary: The cited D-body paper contrasts animal cytoplasmic miRNA processing with plant nuclear processing and does not directly support HEN1 cytoplasmic localization.
Reason: HEN1 may influence cytoplasmic small-RNA turnover indirectly, but PMID:17442570 does not provide direct HEN1 cytoplasm evidence.
GO:0008173 RNA methyltransferase activity
IDA
PMID:15705854
Methylation as a crucial step in plant microRNA biogenesis.
MODIFY
Summary: Experimental Arabidopsis work supports HEN1-dependent methylation/protection of miRNAs and siRNAs. The HEN1 crystal structure (PDB 3HTX) confirms it is a SAM-dependent RNA methyltransferase, with the AdoHcy (SAH) product bound in a Rossmann MTase domain as in classical AdoMet-dependent MTases.
Reason: Use the small-RNA terminal 2-prime-O-methyltransferase term to capture HEN1 substrate and reaction specificity.
Supporting Evidence:
file:ARATH/HEN1/HEN1-deep-research-falcon.md
HEN1 catalyzes transfer of a methyl group from AdoMet to the 2-prime-OH of the 3-prime-terminal nucleotide of small RNAs.
file:ARATH/HEN1/HEN1-uniprot.txt
Methyltransferase that adds a methyl group to the ribose of the last nucleotide of small RNAs (sRNAs).
PMID:19812675
A conserved S-adenosyl-l-methionine-dependent RNA methyltransferase, HUA ENHANCER 1 (HEN1), and its homologues are responsible for this specific modification.
PMID:19812675
The MTase domain of HEN1 adopts a core Ξ±/Ξ² Rossmann structure, in which the cofactor product AdoHcy is bound as in classical S-adenosyl-l-methionine (AdoMet)-dependent MTases
GO:0008173 RNA methyltransferase activity
IMP
PMID:16111943
Methylation protects miRNAs and siRNAs from a 3'-end uridyla...
MODIFY
Summary: Experimental Arabidopsis work supports HEN1-dependent methylation/protection of miRNAs and siRNAs. The HEN1 crystal structure (PDB 3HTX) shows a catalytic Mg2+ ion coordinated by the 2-prime and 3-prime hydroxyls of the 3-prime-terminal nucleotide and four invariant active-site residues, defining a Mg2+-dependent 2-prime-O-methylation mechanism.
Reason: Use the small-RNA terminal 2-prime-O-methyltransferase term to capture HEN1 substrate and reaction specificity.
Supporting Evidence:
file:ARATH/HEN1/HEN1-deep-research-falcon.md
HEN1 catalyzes transfer of a methyl group from AdoMet to the 2-prime-OH of the 3-prime-terminal nucleotide of small RNAs.
file:ARATH/HEN1/HEN1-uniprot.txt
Methyltransferase that adds a methyl group to the ribose of the last nucleotide of small RNAs (sRNAs).
PMID:19812675
Metal ion coordination by both 2' and 3' hydroxyls on the 3'-terminal nucleotide and four invariant residues in the active site of the methyltransferase domain suggests a novel Mg(2+)-dependent 2'-O-methylation mechanism.
PMID:19812675
both the 2β€² and 3β€² hydroxyls of G22m and the side chains of four invariant residues (E796, E799, H800 and H860) are coordinated to a metal ion, Mg2+
GO:0035279 miRNA-mediated gene silencing by mRNA destabilization
IMP
PMID:15851028
microRNA-directed phasing during trans-acting siRNA biogenes...
KEEP AS NON CORE
Summary: HEN1 supports miRNA-mediated gene silencing by mRNA destabilization by methylating and stabilizing the small-RNA duplexes that feed these pathways.
Reason: HEN1 is required for stable miRNA/siRNA pathway outputs, but the direct catalytic step is small-RNA 2-prime-O-methylation rather than Dicer cleavage or AGO-mediated target repression.
Supporting Evidence:
file:ARATH/HEN1/HEN1-uniprot.txt
Can methylate 3'-end of microRNAs (miRNAs), small interfering RNAs (siRNas) and trans-acting small interfering RNAs (ta-siRNAs).
GO:0035196 miRNA processing
IMP
PMID:12747833
Arabidopsis HEN1: a genetic link between endogenous miRNA co...
ACCEPT
Summary: HEN1 supports miRNA processing by methylating and stabilizing the small-RNA duplexes that feed these pathways.
Reason: HEN1 methylation is a required post-dicing maturation/protection step in plant miRNA biogenesis, so miRNA processing is supported even though the more specific catalytic role is small-RNA 2-prime-O-methylation.
Supporting Evidence:
file:ARATH/HEN1/HEN1-uniprot.txt
Can methylate 3'-end of microRNAs (miRNAs), small interfering RNAs (siRNas) and trans-acting small interfering RNAs (ta-siRNAs).
file:ARATH/HEN1/HEN1-deep-research-falcon.md
HEN1 methylation is a defining post-dicing step in plants that stabilizes miRNAs and siRNAs.
PMID:19812675
A subset of small RNAs, such as microRNAs and small interfering RNAs (siRNAs) in plants, Piwi-interacting RNAs in animals and siRNAs in Drosophila, requires an additional crucial step for their maturation; that is, 2'-O-methylation on the 3' terminal nucleotide.
GO:0009909 regulation of flower development
IMP
PMID:11917084
Formation of corymb-like inflorescences due to delay in bolt...
KEEP AS NON CORE
Summary: hen1 mutants affect regulation of flower development.
Reason: The developmental process is a downstream phenotype of altered miRNA/siRNA stability rather than a direct HEN1 function.
GO:0010093 specification of floral organ identity
IMP
PMID:11874905
HEN1 functions pleiotropically in Arabidopsis development an...
KEEP AS NON CORE
Summary: hen1 mutants affect specification of floral organ identity.
Reason: The developmental process is a downstream phenotype of altered miRNA/siRNA stability rather than a direct HEN1 function.

Core Functions

Small-RNA terminal 2-prime-O-methyltransferase activity. HEN1 transfers a methyl group from SAM to the 2-prime-OH of the 3-prime-terminal ribose of miRNA/miRNA-star and siRNA/siRNA-star duplexes.

Supporting Evidence:
  • file:ARATH/HEN1/HEN1-deep-research-falcon.md
    HEN1 is a small RNA 3-prime-terminal 2-prime-O-methyltransferase.
  • file:ARATH/HEN1/HEN1-uniprot.txt
    Methyltransferase that adds a methyl group to the ribose of the last nucleotide of small RNAs (sRNAs).
  • file:interpro/panther/PTHR21404/PTHR21404-entries.csv
    Q9C5Q8,Small RNA 2'-O-methyltransferase

Protection and stabilization of plant small RNAs. HEN1-installed methylation protects miRNAs and siRNAs from HESO1/URT1-dependent uridylation, trimming, and degradation, preserving RNA-silencing competence.

Supporting Evidence:
  • file:ARATH/HEN1/HEN1-deep-research-falcon.md
    HEN1 methylation is a defining post-dicing step in plants that stabilizes miRNAs and siRNAs.
  • file:ARATH/HEN1/HEN1-uniprot.txt
    This protects the 3'-end of sRNAs from uridylation activity and subsequent degradation.

References

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Suggested Questions for Experts

Q: Should piRNA-processing IBA transfer be excluded systematically for plant HEN1 proteins without piRNA pathway evidence?

Q: Should GO add a process term such as small RNA 3-prime-end protection by 2-prime-O-methylation for HEN1-dependent protection from uridylation, trimming, and decay?

Suggested Experiments

Experiment: Quantify methylation efficiency of Arabidopsis HEN1 on defined miRNA, siRNA, and ta-siRNA duplex substrates with varying overhang geometry.

Experiment: Map the compartment and timing of HEN1 methylation relative to DCL1/HYL1 processing and AGO1 loading using endogenous tagged proteins.

Deep Research

Falcon

(HEN1-deep-research-falcon.md)

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