Equisetopsida - Morphology

Morphology

The Sphenophytes comprise photosynthesising, "segmented", hollow stems, sometimes filled with pith. At the junction ("node", see diagram) between each segment is a whorl of leaves. In the only extant genus Equisetum, these are small leaves (microphylls) with a singular vascular trace. However, sphenophyte leaves probably arose by the reduction of a megaphyll, as evidenced by early fossil forms such as Sphenophyllum, in which the leaves are broad with branching veins. The plumbing of these leaves is interesting: the vascular traces trifurcate at the junctions, with one thread going to the microphyll, and the other two moving left and right to merge with the new branches of their neighbours. The vascular system itself curiously resembles that of the vascular plants' eustele, which evolved convergently. A primary xylem contains carinal canals; in the Calamitaceae, secondary xylem (but not secondary phloem) can be secreted as the cambium grows outwards, producing a woody stem, and allowing the plants to grow as high as 10m. The cortex itself contains valecular canals; due to the softer nature of the phloem, these are very rarely seen in fossil instances.

The plant does not bear a coherent root system but underground rhizomes, from which roots and aerial axes emerge.

The plant contains an intercalary meristem: that is to say, each segment of the stem grows as the plant gets taller. This contrasts with the seed plants, which contain an apical meristem - i.e. new growth comes only from growing tips (and widening of stems). Growth was determinate - i.e. the plants' phenotype dictated a maximum height, which the plant would grow to then get no higher.

Sphenophytes bear cones (technically strobili, sing. strobilus) at the tips of some stems. These cones comprise spirally arranged sporophores, which bear spores in four clusters, and in extant sphenophytes cover the spores externally - like four sacs hanging from an umbrella, with its handle embedded in the central cone body. In extinct groups, further protection was afforded to the spores by the presence of whorls of bracts - big pointy microphylls protruding from the cone.

The spores themselves bear characteristic elaters, distinctive spring-like attachments which are hygroscopic: i.e. they change their configuration in the presence of water, helping the spores move and aiding their dispersal. Dispersal is aided in the first instance by laterally dehiscing sporangia, which pop open and scatter spores.

The extant horsetails are mostly homosporous, but this is conspicuously not the case in the past.