module: dmel_cytoplasmic_dynein_dynactin
title: Cytoplasmic dynein and dynactin in Drosophila melanogaster
label: Cytoplasmic dynein and dynactin
module_type: PROTEIN_COMPLEX
description: >-
  Cytoplasmic dyneins are multi-subunit minus-end-directed microtubule motors.
  Cytoplasmic dynein 1, built around the heavy chain Dhc64C, moves organelles,
  vesicles, mRNAs and chromosomes toward microtubule minus ends and acts in spindle
  assembly, nuclear positioning and oocyte polarity. It contains intermediate chains
  (Short wing, sw), light intermediate chains (Dlic) and LC8, Tctex and roadblock
  families of light chains. Dynein 1 is processive only when bound to dynactin and a
  cargo adaptor. Dynactin is a large complex with a short filament of Arp1 capped at
  the barbed end by capping protein and at the pointed end by a complex including
  Arp10 (Arp11), plus a projecting shoulder and arm formed by p150Glued
  (DCTN1-p150), p50 dynamitin (DCTN2-p50) and p24 (DCTN3-p24). Cytoplasmic dynein 2,
  with the heavy chain Beethoven (btv), drives retrograde intraflagellar transport
  in the cilia of sensory neurons.
summary: >-
  Cytoplasmic dynein 1 and dynein 2 motors with their light chains, and the
  dynactin complex that activates dynein 1.
concepts:
  - {id: GO:0005868, label: cytoplasmic dynein complex}
  - {id: GO:0005869, label: dynactin complex}
parts:
  - id: cytoplasmic_dynein_1
    label: Cytoplasmic dynein 1
    role: minus-end-directed motor for cytoplasmic transport
    complex: false
    parts:
      - id: dynein_1_heavy_chain
        label: Dynein 1 heavy chain
        role: motor subunit
        genes: [Dhc64C]
        function: {id: GO:0008569, label: minus-end-directed microtubule motor activity}
      - id: dynein_1_intermediate_chain
        label: Dynein 1 intermediate chain
        role: light chain and dynactin binding
        genes: [sw]
      - id: dynein_1_light_intermediate_chain
        label: Dynein 1 light intermediate chain
        role: cargo adaptor binding
        genes: [Dlic]
  - id: cytoplasmic_dynein_2
    label: Cytoplasmic dynein 2
    role: retrograde intraflagellar transport motor
    complex: false
    parts:
      - id: dynein_2_heavy_chain
        label: Dynein 2 heavy chain
        role: motor subunit
        genes: [btv]
        function: {id: GO:0008569, label: minus-end-directed microtubule motor activity}
        processes:
          - {id: GO:0042073, label: intraciliary transport}
      - id: dynein_2_light_intermediate_chain
        label: Dynein 2 light intermediate chain
        role: dynein 2 light intermediate chain
        genes: [CG3769]
  - id: dynein_light_chains
    label: Cytoplasmic dynein light chains
    role: light chains bound to the intermediate chains
    description: >-
      LC8-family (ctp, Cdlc2), Tctex-family (Dlc90F, CG7276) and roadblock-family
      (robl and its paralogs robl22E, robl37BC, robl62A) light chains, plus the
      uncharacterized FlyBase member CG5359. Which light chains associate with dynein
      1 or dynein 2, and in which tissues the paralogs act, is not established for all
      of them.
    genes: [ctp, Cdlc2, Dlc90F, CG7276, robl, robl22E, robl37BC, robl62A, CG5359]
    complex: false
  - id: dynactin
    label: Dynactin complex
    role: dynein 1 activator and cargo linker
    parts:
      - id: dynactin_projection
        label: Dynactin shoulder and projection arm
        role: dynein and microtubule binding
        description: >-
          p150Glued (DCTN1-p150), which binds dynein intermediate chains and
          microtubules, with p50 dynamitin (DCTN2-p50) and p24 (DCTN3-p24).
        genes: [DCTN1-p150, DCTN2-p50, DCTN3-p24]
        complex_term: {id: GO:0005869, label: dynactin complex}
      - id: dynactin_arp1_filament
        label: Arp1 filament and its end complexes
        role: cargo-binding backbone
        description: >-
          The Arp1 filament with the barbed-end capping protein (cpa, cpb), and the
          pointed-end subunits Arp10 (Arp11), DCTN4-p62, DCTN5-p25 and DCTN6-p27.
        genes: [Arp1, cpa, cpb, Arp10, DCTN4-p62, DCTN5-p25, DCTN6-p27]
        complex_term: {id: GO:0005869, label: dynactin complex}
      - id: dynactin_dred
        label: Dred (DCTN1-p150 paralog)
        role: DCTN1-p150 paralog of unknown role
        genes: [Dred]
        complex: false
connections:
  - source: dynactin
    target: cytoplasmic_dynein_1
    connection_type: POSITIVELY_REGULATES
    description: >-
      Dynactin, together with a cargo adaptor, activates processive minus-end
      movement of dynein 1.
notes: >-
  Boundary. The FlyBase dynein light chain group (DYN-LC) also contains two axonemal
  light chains, ODA-Dnal1 (outer-arm LC1 homolog) and Dnali1 (inner-arm p28
  homolog); they are excluded here and modelled in dmel_axonemal_dynein_arms. Cargo
  adaptors (for example BicD), Lis1 and NudE are regulators and not parts. The
  dynein 2 intermediate chains and dedicated light chains are not in the FlyBase
  groups and are not modelled. cpa and cpb also form the stand-alone actin capping
  protein (dmel_f_actin_capping_protein).
groups: [DYNA-C, DYNC, DYN-LC, DYN-LIC, DYNC-IC]
exclude: [ODA-Dnal1, Dnali1]
