Lycopodium root meristem dynamics supports homology between shoots and roots in lycophytes

Roots have played a pivotal role in the conquest of land by vascular plants, yet their origin has remained enigmatic. Palaeobotanical evidence suggests that roots may have originated from subterranean shoots in some lycophyte species. If this hypothesis is correct, it would follow that the roots and shoots of extant lycophytes share fundamental developmental mechanisms. We tracked meristem dynamics in root and shoot apices of Lycopodium clavatum using a thymidine analogue and expression patterns of histone H4, respectively. Then we compared the meristem dynamics of roots and shoots to identify developmental similarities. Both apical meristems contained a quiescent tissue characterised by a low frequency of cell division. Actively dividing cells appeared in the quiescent tissue during dichotomous branching of both roots and shoots. As a result, the parental meristem divides into two daughter meristems, which give rise to new root or shoot apices. These striking similarities in meristem dynamics provide new neobotanical data that support the shoot-origin hypothesis of lycophyte roots. Although Lycopodium roots may have originated from subterranean shoots of Devonian lycophytes, these shoots may have changed into root-bearing axes in other extant lycophyte lineages.