The genus Equisetum is unique in its mode of branching. The branches are borne at the nodes, but they develop exogenously from the top of the nodal ring, alternating with the sheath segments or leaves. The branch buds develop between leaves rather than being axillary to the leaf, which is the arrangement found in seed plants, or with no relation to the leaves, as in ferns. It is obvious that the branches are associated with the node below them, rather than the nQde above, because if the number of leaves at a node changes, the number of branches is the same as the number of leaves below it. Although exogenous in origin, the branch bud is enclosed by the leaf sheath and must grow out through its base. Where it emerges, a fringe of sheath tissue is left around the branch base as a collar. The base of the branch is surrounded by a sheath-like structure, which differs from the regular nodal sheaths in being usually scarious, brownish in color, and unvascularized, and in lacking stomata and a subjacent internode. This structure was described at length by Duval-Jouve (1864, as gaine basilaire) who followed Vaucher (1821) in considering it a protective structure for the branch bud, analogous to the bud scales of phanerogams. Milde (1867, p. 155) thoroughly discussed the Asthiille (Ochreola). He pointed out that it is always different in color from the branch sheaths, with short, blunt teeth. According to Milde, the ochreole does not grow from the branch but rather develops from the inner cylinder of the stem. He claimed it lacked commissures so was not of fused leaves, but of a single, toothed leaf. The outer (abaxial) angle is different from the others in being thick, with a vascular bundle, chlorophyllous cells, and scattered stomata. Thus the ochreola represents a bract with a midrib, in the axil of which the branch develops. Milde emphasized that the ochreole does not terminate an internode, as do the branch and stem sheaths. It belongs not to an internode, but rather to the whole branch. He also called attention to the similarity between this and the sheath subtending the juvenile plant as it develops from the embryo. On a later page in the same publication Milde (1867, p. 379) corrected and expanded his earlier description. The Asthiille, unlike a bract, belongs to the branch rather than the main axis, and is the first leaf whorl on the branch. He compared it to the first leaf (the prophyll) on most monocot branches. Janczewski (1876) described the initiating bud as forming a vegetative cone and a basal portion. As the vegetative cone develops, the upper part of the basal portion is elevated to form a rim around the vegetative cone, and this rim represents the first sheath of the branch. Janczewski found Milde's interpretation of this as a bract subtending the branch very odd (bien singuliere). DeBlock (1923) described the ochreole as membranous and becoming brown, with no trace of stomata, chlorophyll, or vascular tissue. Johnson (1937) in describing hydathodes in Equisetum leaves, commented on their absence on the ochreole, which is thin and usually devoid of vascular tissue. Even when vasculated, the ochreole
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