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First published online 16 June 2004
doi: 10.1242/dev.01220
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1 Laboratory for Vertebrate Body Plan, Center for Developmental Biology (CDB),
RIKEN Kobe, 2-2-3 Minatojima-minamimachi, Chuo-ku, Kobe 650-0046, Japan
2 Laboratory for Animal Resources and Genetic Engineering, Center for
Developmental Biology (CDB), RIKEN Kobe, 2-2-3 Minatojima-minamimachi,
Chuo-ku, Kobe 650-0046, Japan
* Author for correspondence (e-mail saizawa{at}cdb.riken.jp)
Accepted 8 April 2004
Otx2 expression in the forebrain and midbrain was found to be
regulated by two distinct enhancers (FM and FM2) located at 75 kb 5'
upstream and 115 kb 3' downstream. The activities of these two enhancers
were absent in anterior neuroectoderm earlier than E8.0; however, at E9.5
their regions of activity spanned the entire mesencephalon and diencephalon
with their caudal limits at the boundary with the metencephalon or isthmus. In
telencephalon, activities were found only in the dorsomedial aspect. Potential
binding sites of OTX and TCF were essential to FM activity, and TCF sites were
also essential to FM2 activity. The FM2 enhancer appears to be unique to
rodent; however, the FM enhancer region is deeply conserved in gnathostomes.
Studies of mutants lacking FM or FM2 enhancer demonstrated that these
enhancers indeed regulate Otx2 expression in forebrain and midbrain.
Development of mesencephalic and diencephalic regions was differentially
regulated in a dose-dependent manner by the cooperation between Otx1
and Otx2 under FM and FM2 enhancers: the more caudal the structure
the higher the OTX dose requirement. At E10.5
Otx1–/–Otx2
FM/FM
mutants, in which Otx2 expression under the FM2 enhancer remained,
exhibited almost complete loss of the entire diencephalon and mesencephalon;
the telencephalon did, however, develop.
Key words: Otx2, Otx1, Enhancer, Forebrain, Midbrain, Anterior neuroectoderm, WNT signaling, Mouse
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