Fig. 7. Summary of the expression patterns of guidance molecules involved in the
zone-to-zone targeting of vomeronasal axons. The upper part depicts
schematically the zone-to-zone projection in the vomeronasal system. Axons
from the apical zone of the VNO terminate in the anterior AOB, whereas basal
axons terminate in the posterior AOB. Note that apical and basal axons enter
the AOB at its medial margin. The lower part summarises the expression
patterns of three families of axon guidance molecules described so far in the
vomeronasal projection. The expression patterns provided in this study suggest
that Robos and Slits act predominantly on basal axons. Robo2 appears to be the
principal guidance receptor for basal axons, whereas Robo1 is uniformly
expressed and later downregulated, leaving only faint expression on both
apical and basal axons (broken line). This suggests a model in which basal
axons expressing Robos more strongly than apical axons navigate to the
posterior AOB due to repulsive interactions, with Slit proteins secreted from
the anterior AOB. Ephrin A proteins and neuropilin 2 instead operate mainly on
the apical subpopulation of VNO axons. Apical axons with higher ephrin A
expression levels than basal axons project to the anterior AOB, with express
higher concentrations of Epha proteins than the posterior AOB. In this scheme,
Epha/ephrin A interactions would guide VNO axons on the basis of an attractive
guidance mechanism, consistent with data from the stripe assay
(Knöll et al., 2001). In
turn, apical axons expressing neuropilin 2 are repelled by class 3
semaphorins, which are uniformly localised in the AOB. Neuropilin 2 in the
anterior AOB might sequester semaphorins, thus rendering apical axons
sensitive to only the semaphorins in the posterior AOB
(Cloutier et al., 2000;
Walz et al., 2002).