Alpha ring (26S proteasome)
alpha_ring_somatic_complex
Participant: Protein Complex: Alpha ring (26S proteasome)
Protein Complex:
26S proteasome in Drosophila melanogaster
The 26S proteasome degrades proteins tagged with polyubiquitin chains. Its 20S core particle is a barrel of four stacked heptameric rings, two outer rings of alpha subunits that gate substrate entry and two inner rings of beta subunits, three of which (beta1, beta2 and beta5) are N-terminal threonine proteases with caspase-like, trypsin-like and chymotrypsin-like specificities. The 19S regulatory particle caps the core. Its base contains six AAA+ ATPases (Rpt1-Rpt6) that unfold substrates and open the core gate, together with the scaffolds Rpn1 and Rpn2. Ubiquitin receptors (Rpn10 and Rpn13) bind ubiquitylated substrates, the lid metalloprotease Rpn11 removes ubiquitin chains before degradation, and Uch-L5 bound to Rpn13 trims chains. Drosophila has a testis-specific 26S proteasome in which paralogs replace several alpha, beta, Rpt, Rpn and Uch-L5 subunits.
proteasome complexGO:0000502
FB:FBgg0000208
FlyBase gene group PSM: Proteasome subunits
FlyBase release FB2026_03 groups 50 D. melanogaster genes as proteasome subunits (PSM); this module models that group.
FB:FBgg0001846
FlyBase gene group PSM-V: Proteasome 26s complex variants
FlyBase release FB2026_03 groups 50 D. melanogaster genes as proteasome 26s complex variants (PSM-V); this module models that group.
Somatic and testis-specific variants follow the FlyBase groups PSM26S and PSM26S-T; subunits shared by both appear in each variant. Prosalpha1R is a member of both FlyBase variant groups. The base, lid and ubiquitin-receptor split follows the conserved eukaryotic 19S organisation; Rpn10 and Rpn13 are modelled separately because their placement between base and lid is not sharply defined. Catalytic activity is asserted only for the canonical beta1 (Prosbeta1), beta2 (Prosbeta2) and beta5 (Prosbeta5) subunits; whether the testis paralogs Prosbeta2R1, Prosbeta2R2, Prosbeta5R1 and Prosbeta5R2 keep catalytic activity is not asserted. Likewise ATPase and deubiquitinase activities are not asserted for the testis paralogs Rpt3R, Rpt4R, Rpt6R and Uch-L5R. FlyBase groups realized by this module: PSM (FBgg0000208), PSM-V (FBgg0001846). Subgroups folded into it: T1P, PSM20S, PSM19S, PSM20SA, PSM20SB, PSM20SB-NC, RPN, RPT, PSM26S-T, PSM26S. Participants were generated from the FlyBase gene-group membership and mapped to UniProtKB by projects/FLYBASE_GENE_GROUPS/generate_modules.py from module_specs/dmel_proteasome.yaml. None of the member genes has a gene review in this repository yet.
18Nodes
7Parts
5Variant Sets
10Variants
14Annotons
1Connections
Derived QC
Recommended-field compliance
100.0%
recommended fields populated
All recommended fields populated.
Module deep research
✗ none found
No MODULE:dmel_proteasome 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
0 gene-function check(s) reflected in core functions · 0 conflict(s) · 16 evidence or coverage gap(s) (16 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.
| Annoton | Participant | Function | Assessment |
|---|---|---|---|
| beta_ring_somatic_complex/prosbeta1 | Prosbeta1 UniProtKB:A0AQH0 Complex Unit |
threonine-type endopeptidase activity GO:0004298 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| beta_ring_somatic_complex/prosbeta2 | Prosbeta2 UniProtKB:Q9VUJ1 Complex Unit |
threonine-type endopeptidase activity GO:0004298 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| beta_ring_somatic_complex/prosbeta5 | Prosbeta5 UniProtKB:A1Z8C3 Complex Unit |
threonine-type endopeptidase activity GO:0004298 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| beta_ring_testis_complex/prosbeta1 | Prosbeta1 UniProtKB:A0AQH0 Complex Unit |
threonine-type endopeptidase activity GO:0004298 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_somatic_complex/rpt1 | Rpt1 UniProtKB:A1Z724 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_somatic_complex/rpt2 | Rpt2 UniProtKB:P48601 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_somatic_complex/rpt3 | Rpt3 UniProtKB:Q9V405 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_somatic_complex/rpt4 | Rpt4 UniProtKB:Q9W414 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_somatic_complex/rpt5 | Rpt5 UniProtKB:Q9V3V6 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_somatic_complex/rpt6 | Rpt6 UniProtKB:O18413 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_testis_complex/rpt1 | Rpt1 UniProtKB:A1Z724 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_testis_complex/rpt2 | Rpt2 UniProtKB:P48601 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| base_testis_complex/rpt5 | Rpt5 UniProtKB:Q9V3V6 Complex Unit |
proteasome-activating activity GO:0036402 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| lid_somatic_complex/rpn11 | Rpn11 UniProtKB:Q9V3H2 Complex Unit |
metal-dependent deubiquitinase activity GO:0140492 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| lid_testis_complex/rpn11 | Rpn11 UniProtKB:Q9V3H2 Complex Unit |
metal-dependent deubiquitinase activity GO:0140492 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| receptors_somatic_uch_l5 | Uch-L5 UniProtKB:Q9XZ61 Gene |
cysteine-type deubiquitinase activity GO:0004843 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
Gene-review completeness (0/50 grounded genes reviewed)
0 complete review(s) · 0 with deep research · 50 missing review · 0 reviewed but lacking deep research
| Gene | Review | Complete | Deep research |
|---|---|---|---|
| Prosbeta1 A0AQH0 | ✗ | — | — |
| Rpt1 A1Z724 | ✗ | — | — |
| Prosbeta5 A1Z8C3 | ✗ | — | — |
| Prosalpha1R E1JGZ9 | ✗ | — | — |
| Rpn9 E2QD34 | ✗ | — | — |
| Rpt6 O18413 | ✗ | — | — |
| Prosalpha6 P12881 | ✗ | — | — |
| Prosalpha3 P18053 | ✗ | — | — |
| Prosalpha4 P22769 | ✗ | — | — |
| Rpn3 P25161 | ✗ | — | — |
| Rpn8 P26270 | ✗ | — | — |
| Prosalpha2 P40301 | ✗ | — | — |
| Prosbeta6 P40304 | ✗ | — | — |
| Rpt2 P48601 | ✗ | — | — |
| Rpn10 P55035 | ✗ | — | — |
| Prosalpha4T1 Q24178 | ✗ | — | — |
| Prosalpha4T2 Q27575 | ✗ | — | — |
| Rpn13 Q7K2G1 | ✗ | — | — |
| Rpn6 Q7KLV9 | ✗ | — | — |
| Prosbeta2R2 Q7KTR5 | ✗ | — | — |
| Prosbeta4R1 Q7KU19 | ✗ | — | — |
| Rpn12R Q7KUL5 | ✗ | — | — |
| Prosbeta5R2 Q8INY4 | ✗ | — | — |
| Prosalpha5 Q95083 | ✗ | — | — |
| Rpn7 Q9V3G7 | ✗ | — | — |
| Rpn11 Q9V3H2 | ✗ | — | — |
| Rpn2 Q9V3P6 | ✗ | — | — |
| Rpt5 Q9V3V6 | ✗ | — | — |
| Rpn5 Q9V3Z4 | ✗ | — | — |
| Rpt3 Q9V405 | ✗ | — | — |
| Rpn12 Q9V436 | ✗ | — | — |
| Prosalpha7 Q9V5C6 | ✗ | — | — |
| Prosalpha3T Q9VA12 | ✗ | — | — |
| Rpt6R Q9VA54 | ✗ | — | — |
| Rpt3R Q9VH79 | ✗ | — | — |
| Prosbeta4 Q9VJJ0 | ✗ | — | — |
| Prosalpha6T Q9VK14 | ✗ | — | — |
| Prosbeta7 Q9VNA5 | ✗ | — | — |
| Prosbeta4R2 Q9VQE5 | ✗ | — | — |
| Rpt4R Q9VTQ9 | ✗ | — | — |
| Prosbeta2 Q9VUJ1 | ✗ | — | — |
| Rpn1 Q9VW54 | ✗ | — | — |
| Uch-L5R Q9VYQ3 | ✗ | — | — |
| Prosbeta5R1 Q9W1S5 | ✗ | — | — |
| Rpt4 Q9W414 | ✗ | — | — |
| Prosbeta2R1 Q9W470 | ✗ | — | — |
| Rpn13R Q9W4E6 | ✗ | — | — |
| Prosbeta3 Q9XYN7 | ✗ | — | — |
| Uch-L5 Q9XZ61 | ✗ | — | — |
| Prosalpha1 Q9XZJ4 | ✗ | — | — |
Details
Context
Drosophila melanogasterNCBITaxon:7227
26S proteasomeProtein Complexdmel_proteasome
Ubiquitin-dependent ATP-driven protease formed by the 20S core particle and 19S regulatory particle, with a testis-specific paralog variant.
proteasome complexGO:0000502
Context
Drosophila melanogasterNCBITaxon:7227
Connections
The regulatory particle unfolds deubiquitylated substrates and translocates them into the core particle.
Part 1: proteolytic chamber
20S core particleProtein Complexcore_particle
proteasome core complexGO:0005839
Part 1: gated entry ring
Alpha ringProtein Complexalpha_ring
Variant set: Somatic or testis alpha ring (One Or More)
Alpha ring (26S proteasome)Protein Complexalpha_ring_somatic
Annotons
Alpha ring (testis-specific 26S proteasome)Protein Complexalpha_ring_testis
Annotons
Alpha ring (testis-specific 26S proteasome)
alpha_ring_testis_complex
Participant: Protein Complex: Alpha ring (testis-specific 26S proteasome)
Protein Complex:
Part 2: catalytic ring
Beta ringProtein Complexbeta_ring
Variant set: Somatic or testis beta ring (One Or More)
Beta ring (26S proteasome)Protein Complexbeta_ring_somatic
Annotons
Beta ring (26S proteasome)
beta_ring_somatic_complex
Participant: Protein Complex: Beta ring (26S proteasome)
Protein Complex:
Beta ring (26S proteasome)GO:0019774
Active units:
Prosbeta1
Participant: Gene Product: Prosbeta1
Gene Product:
Prosbeta1UniProtKB:A0AQH0
Function:
threonine-type endopeptidase activityGO:0004298
Prosbeta2
Participant: Gene Product: Prosbeta2
Gene Product:
Prosbeta2UniProtKB:Q9VUJ1
Function:
threonine-type endopeptidase activityGO:0004298
Prosbeta5
Participant: Gene Product: Prosbeta5
Gene Product:
Prosbeta5UniProtKB:A1Z8C3
Function:
threonine-type endopeptidase activityGO:0004298
Beta ring (testis-specific 26S proteasome)Protein Complexbeta_ring_testis
Annotons
Beta ring (testis-specific 26S proteasome)
beta_ring_testis_complex
Participant: Protein Complex: Beta ring (testis-specific 26S proteasome)
Protein Complex:
Beta ring (testis-specific 26S proteasome)GO:0019774
Active units:
Prosbeta1
Participant: Gene Product: Prosbeta1
Gene Product:
Prosbeta1UniProtKB:A0AQH0
Function:
threonine-type endopeptidase activityGO:0004298
Part 2: substrate recognition, deubiquitylation and unfolding
19S regulatory particleProtein Complexregulatory_particle
proteasome accessory complexGO:0022624
Part 1: AAA+ ATPase ring and scaffolds
Regulatory particle baseProtein Complexrp_base
Variant set: Somatic or testis base (One Or More)
Base (26S proteasome)Protein Complexbase_somatic
Annotons
Base (26S proteasome)
base_somatic_complex
Participant: Protein Complex: Base (26S proteasome)
Protein Complex:
Base (26S proteasome)GO:0008540
Active units:
Rpt1
Participant: Gene Product: Rpt1
Gene Product:
Rpt1UniProtKB:A1Z724
Function:
proteasome-activating activityGO:0036402
Rpt2
Participant: Gene Product: Rpt2
Gene Product:
Rpt2UniProtKB:P48601
Function:
proteasome-activating activityGO:0036402
Rpt3
Participant: Gene Product: Rpt3
Gene Product:
Rpt3UniProtKB:Q9V405
Function:
proteasome-activating activityGO:0036402
Rpt4
Participant: Gene Product: Rpt4
Gene Product:
Rpt4UniProtKB:Q9W414
Function:
proteasome-activating activityGO:0036402
Rpt5
Participant: Gene Product: Rpt5
Gene Product:
Rpt5UniProtKB:Q9V3V6
Function:
proteasome-activating activityGO:0036402
Rpt6
Participant: Gene Product: Rpt6
Gene Product:
Rpt6UniProtKB:O18413
Function:
proteasome-activating activityGO:0036402
Base (testis-specific 26S proteasome)Protein Complexbase_testis
Annotons
Base (testis-specific 26S proteasome)
base_testis_complex
Participant: Protein Complex: Base (testis-specific 26S proteasome)
Protein Complex:
Base (testis-specific 26S proteasome)GO:0008540
Active units:
Rpt1
Participant: Gene Product: Rpt1
Gene Product:
Rpt1UniProtKB:A1Z724
Function:
proteasome-activating activityGO:0036402
Rpt2
Participant: Gene Product: Rpt2
Gene Product:
Rpt2UniProtKB:P48601
Function:
proteasome-activating activityGO:0036402
Rpt5
Participant: Gene Product: Rpt5
Gene Product:
Rpt5UniProtKB:Q9V3V6
Function:
proteasome-activating activityGO:0036402
Part 2: deubiquitylating lid
Regulatory particle lidProtein Complexrp_lid
Variant set: Rpn12 or Rpn12R lid (One Or More)
Lid (26S proteasome)Protein Complexlid_somatic
Annotons
Lid (26S proteasome)
lid_somatic_complex
Participant: Protein Complex: Lid (26S proteasome)
Protein Complex:
Lid (26S proteasome)GO:0008541
Active units:
Rpn11
Participant: Gene Product: Rpn11
Gene Product:
Rpn11UniProtKB:Q9V3H2
Function:
metal-dependent deubiquitinase activityGO:0140492
Lid (testis-specific 26S proteasome)Protein Complexlid_testis
Annotons
Lid (testis-specific 26S proteasome)
lid_testis_complex
Participant: Protein Complex: Lid (testis-specific 26S proteasome)
Protein Complex:
Lid (testis-specific 26S proteasome)GO:0008541
Active units:
Rpn11
Participant: Gene Product: Rpn11
Gene Product:
Rpn11UniProtKB:Q9V3H2
Function:
metal-dependent deubiquitinase activityGO:0140492
Part 3: ubiquitin chain recognition and trimming
Ubiquitin receptors and Uch-L5Protein Complexubiquitin_receptors
Variant set: Somatic or testis receptor set (One Or More)
Rpn10, Rpn13 and Uch-L5Protein Complexreceptors_somatic
Annotons
Rpn10 Rpn10, Rpn13 and Uch-L5
receptors_somatic_rpn10
Participant: Gene Product: Rpn10
Gene Product:
Rpn10UniProtKB:P55035
Rpn13 Rpn10, Rpn13 and Uch-L5
receptors_somatic_rpn13
Participant: Gene Product: Rpn13
Gene Product:
Rpn13UniProtKB:Q7K2G1
Uch-L5 Rpn10, Rpn13 and Uch-L5
receptors_somatic_uch_l5
Participant: Gene Product: Uch-L5
Gene Product:
Uch-L5UniProtKB:Q9XZ61
Function
cysteine-type deubiquitinase activityGO:0004843
Rpn10, Rpn13R and Uch-L5RProtein Complexreceptors_testis
Annotons
Rpn10 Rpn10, Rpn13R and Uch-L5R
receptors_testis_rpn10
Participant: Gene Product: Rpn10
Gene Product:
Rpn10UniProtKB:P55035
Rpn13R Rpn10, Rpn13R and Uch-L5R
receptors_testis_rpn13r
Participant: Gene Product: Rpn13R
Gene Product:
Rpn13RUniProtKB:Q9W4E6
Uch-L5R Rpn10, Rpn13R and Uch-L5R
receptors_testis_uch_l5r
Participant: Gene Product: Uch-L5R
Gene Product:
Uch-L5RUniProtKB:Q9VYQ3