Table 1.
Examples of molecules involved in dendrite morphogenesis
| Molecules | Functions in dendrite morphogenesis | Refs |
|---|---|---|
| Transcription regulators | ||
| CUT | A homeodomain protein that functions as a multi-level regulator of type-specific dendrite morphology of larval dendritic arborization neurons | 27 |
| Abrupt | A BTB–zinc finger protein that is expressed in larval class I dendritic arborization neurons, where it limits dendrite size and branching complexity | 28,29 |
| Collier (also known as Knot) | A member of the early B cell factor or olfactory neuronal transcription factor 1 family that functions in combination with CUT to distinguish class IV from class III dendritic arborization neuron dendrites | 30–32 |
| Spineless | A fly homologue of the mammalian dioxin receptor that diversifies dendrite morphology of larval dendritic arborization neurons | 36 |
| ACJ6 and Drifter | POU-domain proteins that regulate dendritic targeting in the fly olfactory system | 39 |
| CREST | A transactivator that regulates Ca2+-dependent dendrite growth of mammalian cortical and hippocampal neurons | 42 |
| NEUROD1 | A basic helix–loop–helix (bHLH) protein that regulates activity-dependent dendrite morphogenesis of mammalian neurons | 41 |
| CREB | A DNA-binding protein that regulates activity-dependent dendrite morphogenesis of mammalian neurons | 43,44 |
| Neurogenin 2 | A bHLH protein that specifies the dendrite morphology of pyramidal neurons in the mammalian neocortex | 40 |
| nBAF complexes | Neuron-specific chromatin remodelling complexes that bind CREST and regulate activity-dependent dendrite development of mammalian CNS neurons; their D. melanogaster homologues are involved in dendrite morphogenesis in dendritic arborization neurons | 33,45 |
| Polycomb group genes | Components of the Polycomb complex that regulate the maintenance of larval dendritic arborization dendrites | 146 |
| Secreted proteins and cell surface receptors | ||
| Neurotrophins and tyrosine kinase receptors | Regulate dendritic growth and complexities of mammalian cortical neurons in a neuronal type-dependent manner | 54,55 |
| BMP7 | Promotes dendrite growth in rat sympathetic neurons | 56 |
| WNT and Dishevelled | Regulate dendrite morphogenesis in mammalian neurons | 72 |
| EPHB1, EPHB2 and EPHB3 | Promote dendritic complexity of mammalian hippocampal neurons | 91 |
| Semaphorins and Plexin–Neuropilin | Regulate dendritic targeting and branching in mammalian cortical neurons and fly olfactory neurons | 48,50,52 |
| Slit and Robo | Regulate dendrite growth and targeting of fly neurons | 46,47,127,128 |
| Netrin and Frazzled | Regulate dendritic targeting of fly motor neurons | 127,128 |
| Reelin | Regulates dendrite growth of mammalian cortical neurons | 135 |
| Cytoskeletal regulators | ||
| RAC and CDC42 | Rho family GTPases that regulate dendrite growth and branching in vertebrate and invertebrate neurons | 59,62,63, 69,71 |
| RHOA | A Rho family GTPase that limits dendrite growth in vertebrate and invertebrate neurons | 60,63,67 |
| Motors | ||
| KIF5 | A kinesin involved in dendrite morphogenesis of mammalian hippocampal neurons | 91 |
| Dynein | Mutations of Dynein cause a distal to proximal shift of the dendrite branching pattern of larval dendritic arborization neurons | 57,58 |
| LIS1 | A dynein-interacting protein linked to lissencephaly; mutations cause reduced dendritic arborization in flies and mice | 57,86–89 |
| Secretory pathway and endocytic pathway | ||
| DAR3 (fly) or SAR1 (mammalian) | A regulator of ER to Golgi transport preferentially required for dendrite growth of mammalian hippocampal neurons and larval dendritic arborization neurons | 82 |
| DAR2 (fly) or SEC23 (mammalian) and DAR6 (fly) or RAB1(mammalian) | Regulators of ER to Golgi transport preferentially required for dendrite versus axonal growth of larval dendritic arborization neurons | 82 |
| RAB5 | A component of the early endocytic pathway that promotes dendrite branching of fly dendritic arborization neurons | 58 |
| Shrub | A fly homologue of SNF7, a component of the ESCRT-III complex, which is a regulator of dendrite branching of larval dendritic arborization neurons | 83 |
| Cell adhesion | ||
| DSCAM (fly) | Mediates self-avoidance of dendrites of the same neuron and enables co-existence of dendrites of different neuron types | 102–105 |
| DSCAM and DSCAML1 (mouse) | Unlike their fly homologues, murine Dscam and Dscaml1 do not undergo extensive alternative splicing; they are required for dendrite self-avoidance but do not confer neuronal identity | 98,109 |
| Hippo pathway | ||
| TRC and FRY | Required for dendritic tiling of larval class IV dendritic arborization neurons | 123 |
| SAX-1 and SAX-2 | Homologues of TRC and FRY required for dendritic tiling in Caenorhabditis elegans | 125 |
| WTS and SAV | Regulators of the maintenance of larval class IV dendritic arborization neurons | 147 |
| Bantam | A microRNA required for dendritic scaling of larval class IV dendritic arborization neurons | 131 |
| Hippo | Regulates dendritic tiling and dendritic maintenance of larval class IV dendritic arborization neurons through the two NDR kinase family members, TRC and WTS, respectively | 147 |
| PIK3–mTOR pathway | ||
| AKT | Promotes dendritic growth and branching of larval class IV dendritic arborization neurons and mammalian cortical neurons | 132–136 |
| PTEN | Limits dendrite growth and branching of larval class IV dendritic arborization neurons | 132 |
| TORC2 | Required for dendritic tiling of larval class IV dendritic arborization neuron | 124 |
| RNA targeting and local translation | ||
| Pumilio and Nanos | RNA-binding proteins required for dendrite growth and branching of larval class III and class IV dendritic arborization neurons | 137 |
| Glorund and Smaug | Bind to Nanos mRNA and regulate dendrite growth of larval class IV dendritic arborization neurons | 138 |
| Staufen 1 | A murine homologue of fly Staufen, which functions in dendritic targeting of ribonucleoprotein particles and dendritic branching | 139 |
| CPEB | A PolyA-binding protein implicated in mRNA trafficking, translational regulation and experience-dependent dendrite growth in frog neurons | 141 |
| FMR1 | Involved in mRNA transport and translation regulation in dendrites, and dendrite morphogenesis | 6 |
| Others | ||
| Origin recognition complex | Regulates dendrite and spine development in mammalian neurons | 80 |
| Anaphase-promoting complex | Specifically regulates axon or dendrite morphogenesis in murine cerebellar granule cells depending on whether the co-activator CDH1 or CDH20 is recruited | 152,153 |
BMP7, bone morphogenetic protein 7 (also known as OP1); CDH, cadherin; CPEB, cytoplasmic polyadenylation element-binding protein; CREB, cAMP response element-binding protein; CREST, Ca2+-responsive transactivator; DSCAM, Down syndrome cell adhesion molecule; DSCAML1, Down syndrome cell adhesion molecule-like protein 1 homologue; EPHB, ephrin receptor type B; ER, endoplasmic reticulum; ESCRT, endosomal sorting complex required for transport; FMR1, fragile X mental retardation 1 protein; FRY, Furry; KIF5, kinesin-related protein 5; mTOR, mammalian target of rapamycin; nBAF, neuron-specific BRG1-associated factor; NDR, nuclear DBF2-related; NEUROD1, neurogenic differentiation factor 1; PI3K, phosphoinositide 3-kinase; POU, PIT1–OCT1–UNC86; PTEN, phosphatase and tensin homologue; RAC, RAS-related C3 botulinum toxin substrate; SAV, Salvador; TORC2, target of rapamycin complex 2; TRC, Tricornered; WTS, Warts.