Satellite phage P4 molecular piracy of helper myovirus P2 morphogenetic, packaging, and lysis machinery

A cross-genome molecular piracy module in which satellite bacteriophage P4 (Enterobacteria phage P4, 11.6 kb, lacking genes for structural capsid, tail, terminase, or lysis proteins) hijacks the morphogenetic, DNA-packaging, and host-lysis apparatus of a coinfecting or lysogenic Peduoviridae helper phage such as Escherichia phage P2 (33.5 kb). The piracy program operates across four tightly coupled stages: (1) Helper prophage derepression, in which P4 derepression protein Epsilon binds the P2 immunity repressor C to prevent operator binding and induce P2 prophage excision and lytic gene expression, while P4 replicates its own 11.6-kb genome autonomously via its bifunctional primase-helicase-origin-binding protein Alpha; (2) Cross-genome late transcriptional transactivation and antitermination, in which the P4 tandem-Ogr-domain zinc-finger protein Delta (paralleling P2's own late activator Ogr) binds the alpha-CTD of host Escherichia coli RNA polymerase to activate P2's four morphogenetic and lysis promoters (PP, PO, PV, PJ), while P4 polarity suppression protein Psu binds and hyper-oligomerizes host transcription termination factor Rho to prevent premature termination across the hijacked P2 late operons; (3) Capsid size redirection, in which P4 protein Sid polymerizes into an external dodecahedral cage around assembling P2 gpN major capsid hexamers and gpO internal protease/scaffold subunits, redirecting procapsid geometry from the helper's T=7 head (60 nm, 35-kb capacity) to a threefold smaller T=4 head (45 nm, 11.6-kb capacity) that physically excludes the full-length P2 helper genome; and (4) Cos-site packaging piracy, in which P4 carries an identical 19-bp 5'-cohesive cos site recognized by the P2 terminase complex (small endonuclease subunit gpM and large ATPase subunit gpP), packaging the 11.6-kb P4 genome through the P2 gpQ portal into the Sid-constricted T=4 capsid before P2 tail attachment and lysis.

MODULE:phage_p4_p2_molecular_piracyCOMPLETECONCRETEBiological Processmodules/phage_p4_p2_molecular_piracy.yaml
release from viral latencyGO:0019046 viral DNA genome replicationGO:0039693 transcription antiterminationGO:0031564 viral capsid assemblyGO:0019069 viral procapsid maturationGO:0046797 viral DNA genome packaging, 5' extended cos packagingGO:0098037
PMID:7643390
The capsid size-determining protein Sid forms an external scaffold on phage P4 procapsids.
Demonstrates that satellite phage P4 protein Sid forms an external dodecahedral scaffold cage around helper P2 gpN/gpO procapsids, constraining assembly to small T=4 heads.
PMID:17931675
Incorporation of scaffolding protein gpO in bacteriophages P2 and P4.
Characterizes the interplay between the P2 internal scaffolding protease gpO and the P4 external scaffold Sid during T=7 versus T=4 procapsid assembly.
PMID:19064277
Functional domains of the bacteriophage P2 scaffolding protein: identification of residues involved in assembly and protease activity.
Defines the N-terminal serine protease domain and C-terminal scaffolding domain of P2 gpO required for both P2 and P4 capsid maturation.
PMID:2193910
A mutation of the transactivation gene of satellite bacteriophage P4 that suppresses the rpoA109 mutation of Escherichia coli.
Establishes that P4 transactivation protein Delta directly interacts with the alpha subunit of Escherichia coli RNA polymerase to activate helper phage P2 late promoters.
PMID:39788982
The Psu protein of phage satellite P4 inhibits transcription termination factor ρ by forced hyper-oligomerization.
Cryo-EM and biochemical demonstration that P4 Psu binds ATP-bound host Rho hexamers and forces inactive helical hyper-oligomerization to suppress transcriptional polarity.
GO:0019069
viral capsid assembly
Defines the assembly of viral capsid subunits and scaffolding factors into procapsids and mature heads.

Exemplar satellite-helper pair: Enterobacteria phage P4 (NCBITaxon:10680) and Bacteriophage P2 (NCBITaxon:2905681). Generalizes across P4-like phage satellites (e.g. Retron-harboring P4 satellites, Phage Inducible Chromosomal Islands / PICIs such as Staphylococcus aureus SaPIs, and ICP1-associated PLEs) that combine helper derepression, late promoter transactivation, external or internal scaffold-mediated capsid remodeling (T=7 to T=4), and packaging site mimicry.

5Nodes
4Parts
0Variant Sets
0Variants
11Annotons
4Connections

Derived QC

Recommended-field compliance

100.0% recommended fields populated

All recommended fields populated.

Module deep research

✗ none found

No MODULE:phage_p4_p2_molecular_piracy deep-research report alongside the module YAML.

Leaf nodes lacking representative members

✓ every leaf node grounds to a representative protein.

Template conformance

✓ every declared conforms_to bundle matches its template motif.

Function consistency and core coverage

11 gene-function check(s) reflected in core functions · 0 conflict(s) · 11 evidence or coverage gap(s) (22 comparison(s))

Retained noncore functions are reported separately; they do not require promotion to core functions. Support from representative genes does not establish conservation across the whole family. Missing reviews or identifiers leave a claim unverified.

AnnotonParticipantFunctionAssessment
bpp4_epsilon_derepression_activity Epsilon (Enterobacteria phage P4)
UniProtKB:P05463
Family Representative
transcription regulator activity
GO:0140110
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp2_c_repressor_activity C (Bacteriophage P2)
UniProtKB:P04132
Family Representative
DNA-binding transcription repressor activity
GO:0001217
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp4_alpha_primase_helicase_activity Alpha (Enterobacteria phage P4)
UniProtKB:P10277
Family Representative
DNA helicase activity
GO:0003678
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp4_delta_transactivator_activity Delta (Enterobacteria phage P4)
UniProtKB:P12551
Family Representative
DNA binding
GO:0003677
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp2_ogr_activator_activity ogr (Bacteriophage P2)
UniProtKB:P08762
Family Representative
DNA-binding transcription activator activity
GO:0001216
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp4_psu_rho_inhibitor_activity psu (Enterobacteria phage P4)
UniProtKB:P05460
Family Representative
enzyme inhibitor activity
GO:0004857
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp4_sid_external_scaffold_activity sid (Enterobacteria phage P4)
UniProtKB:P05461
Family Representative
structural constituent of virion
GO:0039660
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp2_o_internal_scaffold_protease_activity O (Bacteriophage P2)
UniProtKB:P25478
Family Representative
serine-type endopeptidase activity
GO:0004252
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp2_n_major_capsid_activity N (Bacteriophage P2)
UniProtKB:P25477
Family Representative
structural constituent of virion
GO:0039660
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp2_m_terminase_endonuclease_activity M (Bacteriophage P2)
UniProtKB:P25476
Family Representative
endonuclease activity
GO:0004519
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp2_p_terminase_atpase_activity P (Bacteriophage P2)
UniProtKB:P25479
Family Representative
ATP hydrolysis activity
GO:0016887
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass).
bpp4_epsilon_derepression_activity Satellite phage P4 derepression protein Epsilon family
Family
transcription regulator activity
GO:0140110
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp2_c_repressor_activity Phage P2 immunity repressor C HTH family
Family
DNA-binding transcription repressor activity
GO:0001217
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp4_alpha_primase_helicase_activity Satellite phage P4 primase-helicase Alpha family
Family
DNA helicase activity
GO:0003678
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp4_delta_transactivator_activity Phage Ogr/Delta C4 zinc-finger transcriptional activator family
Family
DNA binding
GO:0003677
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp2_ogr_activator_activity Phage Ogr/Delta C4 zinc-finger transcriptional activator family
Family
DNA-binding transcription activator activity
GO:0001216
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp4_psu_rho_inhibitor_activity Satellite phage P4 polarity suppression protein Psu family
Family
enzyme inhibitor activity
GO:0004857
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp4_sid_external_scaffold_activity Satellite phage P4 external scaffolding protein Sid family
Family
structural constituent of virion
GO:0039660
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp2_o_internal_scaffold_protease_activity Phage P2 capsid scaffolding protease gpO family
Family
serine-type endopeptidase activity
GO:0004252
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp2_n_major_capsid_activity Phage P2 major capsid protein gpN family
Family
structural constituent of virion
GO:0039660
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp2_m_terminase_endonuclease_activity Phage P2 terminase small endonuclease subunit gpM family
Family
endonuclease activity
GO:0004519
Family Id Missing
The family has no identifier linking it to a curated family review.
bpp2_p_terminase_atpase_activity Phage P2 terminase large ATPase subunit gpP family
Family
ATP hydrolysis activity
GO:0016887
Family Id Missing
The family has no identifier linking it to a curated family review.

Gene-review completeness (11/11 grounded genes reviewed)

11 complete review(s) · 0 with deep research · 0 missing review · 11 reviewed but lacking deep research

Gene Review Complete Deep research
Alpha P10277 ✓ ✓ ✗
C P04132 ✓ ✓ ✗
Delta P12551 ✓ ✓ ✗
Epsilon P05463 ✓ ✓ ✗
M P25476 ✓ ✓ ✗
N P25477 ✓ ✓ ✗
O P25478 ✓ ✓ ✗
ogr P08762 ✓ ✓ ✗
P P25479 ✓ ✓ ✗
psu P05460 ✓ ✓ ✗
sid P05461 ✓ ✓ ✗

Details

Context
CaudoviricetesNCBITaxon:2731619 Enterobacteria phage P4NCBITaxon:10680 Bacteriophage P2NCBITaxon:2905681
Satellite phage P4 molecular piracy of helper myovirus P2 morphogenesis and packagingBiological Processphage_p4_p2_molecular_piracy
release from viral latencyGO:0019046 viral DNA genome replicationGO:0039693 transcription antiterminationGO:0031564 viral capsid assemblyGO:0019069 viral procapsid maturationGO:0046797 viral DNA genome packaging, 5' extended cos packagingGO:0098037
Context
CaudoviricetesNCBITaxon:2731619 Enterobacteria phage P4NCBITaxon:10680 Bacteriophage P2NCBITaxon:2905681

Connections

Part 1: helper prophage derepression and autonomous satellite replication
Epsilon-mediated derepression of P2 repressor C and Alpha-catalyzed P4 replicationRegulatory Stephelper_derepression_and_satellite_replication_step

Annotons

Satellite P4 derepression protein Epsilon
bpp4_epsilon_derepression_activity
Participant: Family: Satellite phage P4 derepression protein Epsilon family
Family:
Satellite phage P4 derepression protein Epsilon family
Representative Members: Epsilon (Enterobacteria phage P4)UniProtKB:P05463

Function

transcription regulator activityGO:0140110

Processes

release from viral latencyGO:0019046 positive regulation of viral transcriptionGO:0050434

Locations

host cell cytoplasmGO:0030430

Directly binds the dimeric helper phage P2 immunity repressor C, forming inactive heteromeric complexes that strip C from the P2 early operator and derepress the resident P2 prophage without requiring host SOS activation.

Helper P2 immunity repressor C
bpp2_c_repressor_activity
Participant: Family: Phage P2 immunity repressor C HTH family
Family:
Phage P2 immunity repressor C HTH family
Representative Members: C (Bacteriophage P2)UniProtKB:P04132

Function

DNA-binding transcription repressor activityGO:0001217

Processes

maintenance of viral latencyGO:0019044 latency-replication decisionGO:0098689

Locations

host cell cytoplasmGO:0030430

Maintains P2 prophage lysogeny by binding operator sites O1 and O2 at the Pe early promoter; targeted and inactivated by P4 Epsilon upon satellite infection.

Satellite P4 bifunctional primase-helicase Alpha
bpp4_alpha_primase_helicase_activity
Participant: Family: Satellite phage P4 primase-helicase Alpha family
Family:
Satellite phage P4 primase-helicase Alpha family
Representative Members: Alpha (Enterobacteria phage P4)UniProtKB:P10277

Function

DNA helicase activityGO:0003678

Processes

viral DNA genome replicationGO:0039693 DNA replication, synthesis of primerGO:0006269

Locations

host cell cytoplasmGO:0030430

Multifunctional 84.8-kDa replication protein combining C-terminal ori/crr DNA recognition, SF3-type 3'-5' DNA helicase activity, and N-terminal TOPRIM/zinc-ribbon primase activity to drive autonomous bidirectional theta replication of the 11.6-kb P4 genome.

Part 2: cross-genome late promoter transactivation and Rho antitermination
Delta/Ogr activation of P2 late promoters and Psu inhibition of host Rho terminationRegulatory Steplate_operon_transactivation_and_antitermination_step

Annotons

Satellite P4 transactivation protein Delta
bpp4_delta_transactivator_activity
Participant: Family: Phage Ogr/Delta C4 zinc-finger transcriptional activator family
Family:
Phage Ogr/Delta C4 zinc-finger transcriptional activator family
Representative Members: Delta (Enterobacteria phage P4)UniProtKB:P12551

Function

DNA bindingGO:0003677

Processes

positive regulation of viral transcriptionGO:0050434

Locations

host cell cytoplasmGO:0030430

Contains two tandem Ogr-like C4 zinc-finger domains that bind upstream of helper phage P2 late promoters (PP, PO, PV, PJ) at -55 and recruit the alpha-CTD of host Escherichia coli RNA polymerase to activate P2 morphogenetic and lysis gene transcription even in the absence of P2 DNA replication.

Helper P2 late control protein Ogr
bpp2_ogr_activator_activity
Participant: Family: Phage Ogr/Delta C4 zinc-finger transcriptional activator family
Family:
Phage Ogr/Delta C4 zinc-finger transcriptional activator family
Representative Members: ogr (Bacteriophage P2)UniProtKB:P08762

Function

DNA-binding transcription activator activityGO:0001216

Processes

positive regulation of DNA-templated transcriptionGO:0045893

Locations

host cell cytoplasmGO:0030430

Helper phage C4 zinc-finger transcriptional activator that binds P2 late promoters and the host RNA polymerase alpha-CTD, amplified after P4 Epsilon-mediated derepression and Delta transactivation.

Satellite P4 polarity suppression protein Psu
bpp4_psu_rho_inhibitor_activity
Participant: Family: Satellite phage P4 polarity suppression protein Psu family
Family:
Satellite phage P4 polarity suppression protein Psu family
Representative Members: psu (Enterobacteria phage P4)UniProtKB:P05460

Function

enzyme inhibitor activityGO:0004857

Processes

transcription antiterminationGO:0031564 positive regulation of viral transcriptionGO:0050434

Locations

host cell cytoplasmGO:0030430

Coiled-coil V-shaped homodimer that binds the C-terminal ATPase domain of host transcription termination factor Rho, inducing open-helical hyper-oligomerization of Rho into an inactive state to prevent premature Rho-dependent termination across P2 late morphogenetic operons.

Part 3: external scaffold-mediated capsid size redirection (T=7 to T=4)
Sid external dodecahedral cage constriction of P2 gpN/gpO procapsids from T=7 to T=4Protein Complexsid_capsid_size_redirection_step

Annotons

Satellite P4 capsid size-determining external scaffold Sid
bpp4_sid_external_scaffold_activity
Participant: Family: Satellite phage P4 external scaffolding protein Sid family
Family:
Satellite phage P4 external scaffolding protein Sid family
Representative Members: sid (Enterobacteria phage P4)UniProtKB:P05461

Function

structural constituent of virionGO:0039660

Processes

viral capsid assemblyGO:0019069

Locations

host cell cytoplasmGO:0030430

Polymerizes into an external dodecahedral cage of 120 subunits around assembling helper P2 gpN major capsid hexamers, making specific contacts at hexamer threefold axes that force higher shell curvature and redirect procapsid assembly from the T=7 helper geometry (415 gpN subunits, 60 nm) to a T=4 satellite geometry (235 gpN subunits, 45 nm) capable of packaging only the 11.6-kb P4 genome.

Helper P2 internal scaffolding protease gpO
bpp2_o_internal_scaffold_protease_activity
Participant: Family: Phage P2 capsid scaffolding protease gpO family
Family:
Phage P2 capsid scaffolding protease gpO family
Representative Members: O (Bacteriophage P2)UniProtKB:P25478

Function

serine-type endopeptidase activityGO:0004252

Processes

viral procapsid maturationGO:0046797 viral capsid assemblyGO:0019069

Locations

host cell cytoplasmGO:0030430

Bifunctional helper protein whose C-terminal scaffolding domain chaperones gpN shell assembly (acting in concert with P4 Sid during T=4 procapsid assembly) while its N-terminal serine protease domain autocleaves gpO into h7 and cleaves the N-terminal 31 residues of gpN to yield mature N* capsid subunits.

Helper P2 major capsid protein gpN
bpp2_n_major_capsid_activity
Participant: Family: Phage P2 major capsid protein gpN family
Family:
Phage P2 major capsid protein gpN family
Representative Members: N (Bacteriophage P2)UniProtKB:P25477

Function

structural constituent of virionGO:0039660

Processes

viral capsid assemblyGO:0019069

Locations

host cell cytoplasmGO:0030430

HK97-fold major capsid precursor cleaved by gpO protease to N*, assembling either into 60-nm T=7 helper heads or, when caged by P4 Sid via the sir loop region, into 45-nm T=4 satellite heads.

Part 4: cos-site mimicry and helper terminase packaging piracy
P2 terminase (gpM/gpP) cleavage of P4 cos sites and translocation into T=4 procapsidsReactioncos_packaging_piracy_step

Annotons

Helper P2 terminase small endonuclease subunit gpM
bpp2_m_terminase_endonuclease_activity
Participant: Family: Phage P2 terminase small endonuclease subunit gpM family
Family:
Phage P2 terminase small endonuclease subunit gpM family
Representative Members: M (Bacteriophage P2)UniProtKB:P25476

Function

endonuclease activityGO:0004519

Processes

viral DNA genome packaging, 5' extended cos packagingGO:0098037

Locations

host cell cytoplasmGO:0030430

Recognizes and cleaves the conserved 55-bp cos region (producing 19-nt 5'-cohesive ends) shared identically between helper P2 and satellite P4 covalently closed circular monomeric DNA substrates.

Helper P2 terminase large ATPase subunit gpP
bpp2_p_terminase_atpase_activity
Participant: Family: Phage P2 terminase large ATPase subunit gpP family
Family:
Phage P2 terminase large ATPase subunit gpP family
Representative Members: P (Bacteriophage P2)UniProtKB:P25479

Function

ATP hydrolysis activityGO:0016887

Processes

viral DNA genome packaging, 5' extended cos packagingGO:0098037

Locations

host cell cytoplasmGO:0030430

Docks at the P2 gpQ portal vertex of the Sid-constricted T=4 procapsid and uses ATP hydrolysis to package the cos-linearized 11.6-kb P4 genome, triggering external Sid scaffold release and capsid expansion.