mortality/aging
• expected number of double homozygotes are born, but all die within 1 hour of birth
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nervous system
• E14.5-born neurons occupy significantly altered positions relative to wild-type neurons; in wild-type these neurons are in layers V and VI, but in mutant cortex, the neurons remain in the intermediate zone
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• less cell proliferation is observed at E14.5 onward compared to wild-type
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• less foliation observed at P0, with loosely packed Purkinje cells observed
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• severely affected, with marked reduction in thickness at P0
• cortical neurons have altered positioning in neocortex
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• stratification of cortical neurons appears disorganized at P0
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• upper-layer neurons are lost in mutants in late development
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• bulb exhibits hypoplasia
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• reduction in proliferating cells is most severe in cortical subventricular zone (only 16.5% of wild-type cell proliferation)
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cellular
• E14.5-born neurons occupy significantly altered positions relative to wild-type neurons; in wild-type these neurons are in layers V and VI, but in mutant cortex, the neurons remain in the intermediate zone
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