Asymmetric division

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All centrosomes are not equivalent!

DNAMicrotubulesIoDPP mRNA

Yeast spindle

orientation

GFP-tubulin

GFP-tubulinDyn1:His3

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Video

Video

MTOC reorientation to the immunological synapse in single positive

mature T-cells

Ash1 mRNA (encoding a transcriptional repressor of MAT switching) is transported

into the daughter in budding yeast

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Locasome transport requires formin-initiated microfilament cables and a myosin V family member (Myo4p)

3’UTR structure is important

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Ash1 mRNA ash1 mutants don’t switch

Proteins are segregated to the bud too

Both the Ist2p protein and mRNA are localized to the bud

Caenorhabditis elegans

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Video

Caenorhabditis elegans development

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GFP::histone fusionFemale Male Pronuclei

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Video

P-granule partitioning

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P-granules fill at the NE

Cytochalasin

Wild-typepartitioning

Nocodazole

GFP::P granules at the movies

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3 Videos

Localization of Par-2::GFP

Video

Par2::GFP is increasingly restricted to the P lineage

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Video

Par mutant phenotypes

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Setting it all up

Mex-5Mex-6Par3/6/aPKC

Pie-1Par-1Par-2P-granules

Pop goes the

weasel!

Xenopus development

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The first cleavages split the embryo on its three axes and

continue to generate a mass of cells

Video

Veg1 binds to VERA on ER

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Gel shift assayDensity gradient centrifugation

METROorganizes

traffic

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Localization of mRNAs in Drosophila

oskarbicoid

Drosophila disconnects mitosis from cytokinesis

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GFP-tau marks microtubules

Video

Pole cells get polar granules

Microinjection rescues the day!

Anterior-Posterior Gradients

Cascades of gene activity

set up the Drosophila body plan

Loss of Bicoid results in no head!

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Wt larval head

bicoid null

Mislocalization of Nanos gives two tails

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Cellularization traps nuclei with different RNA populations that are translated to

make create different microenvironments

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Bicoid, even-skipped and caudal

Partitioning of Cdx2in the mouse embryo

Deb K, Sivaguru M, Yong HY, Roberts RM. Cdx2 gene expression and trophectoderm lineage specification

in mouse embryos.Science. 2006 Feb 17;311(5763):992-6.

Another Drosophila example: neurogenesis

Muscle precursors Postmitotic Neurons Sensory precursors

Apical surface

Notch

GMC

Cell division in Drosophila epidermoblasts

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Basal

Apical

Video

Neuroblast spindle rotation in metaphase

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Basal

Apical

Video

Asymmetric neuroblast division

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Basal

ApicalVideo

Numb partitioning

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DNA and Numb protein and centrosomes

Anaphase Telophase

Neuroblast

GMC

Apical

Basal

GFP-PON (partner of Numb)

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basal

apicalVideo

PON localization precedes spindle rotation

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Cell expressing both GFP::PON and GFP::tubulin

Video

Asymmetric componentsInscuteable + Partner of Inscuteable (PINS)

Baz(Par3)/Par6/aPKCand Galphai (G-protein)

NumbPartner of Numb (PON)

Neuroblast(Apical)

(Basal)GMC

MyoVI

MirandaStauffen (RNA binding)+ Prospero ( homeodomain

transcr. factor)

+

-

Inscuteable RNAi disrupts rotation

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Note cell still appears to divide asymmetrically

Basal

Apical

Video

Mutations in the tumor suppressor Discs Large cause loss of polarioty

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Wild type Mutant

basal

apical