“The bundle-system of Calamites bears a general resemblance to that of Equisetum. A single leaf-trace enters the stem from each leaf, and passes vertically downwards to the next node. In the simplest cases the bundle here forks, its two branches attaching themselves to the alternating bundles which enter the stem at this node. In other cases both the forks attach themselves to the same bundle, so that, in this case, there is no regular alternation. In other cases, again, the bundle runs past one node without forking, and ultimately forms a junction with the traces of the second node below its starting-point. These variations may all occur in the same specimen. The xylem at the node usually forms a continuous ring, for where the regular dichotomous forks of the bundles are absent their place is usually taken by anastomoses[607].”
As in Equisetum, the xylem at the nodes possesses certain characteristic features which distinguish it from the internodal strands. It has already been pointed out that the xylem of Equisetum increases in breadth at the nodes (p. 251, fig. 55, 4); the same is true of Calamites. In fig. 72, C, we have part of a radial section of a Calamite twig in which the broad mass of short nodal tracheids is clearly shown; this nodal wood forms a prominent projection towards the pith. In the lower part of the section the remains of some spiral protoxylem tracheids are seen in a carinal canal.
Fig. 72.
- A. External xylem elements and cambium, c, with imperfect phloem, × 100.
- B. Carinal canal containing protoxylem, px. × 65.
- C. Radial longitudinal section through nodal xylem, px. × 35.
- D. Phloem elements; s, sieve-tubes; p, p, parenchymatous cells.
- (A–C. After Williamson and Scott. D. After Renault.)
The tracheids of the nodal wood are often reticularly pitted, and so differ in appearance from the ordinary scalariform elements.
It is rare to find the phloem clearly preserved, but in specimens where it has been possible to examine this portion of the vascular bundles, it is found to consist of elongated cambiform cells and sieve-tubes. An unusually perfect specimen has been described by Renault[608] in which the phloem elements are preserved in silica. Fig. 72, D, is copied from one of Renault’s drawings, the sieve-tubes, s, s, show several distinct sieve-plates on the lateral walls of the tubes, reminding one to some extent of the sieve-tubes in a Bracken Fern. The cells, p, p, associated with the sieve-tubes are square-ended elongated parenchymatous elements. Another characteristic feature illustrated by longitudinal sections is the nodal diaphragm; except in the smallest branches the interior of each internode is hollow, and the ring of vascular bundles is separated from the pith-cavity by a band of parenchymatous tissue. At each node this parenchyma extends across the central cavity in the form of a nodal diaphragm, as in the stem of Equisetum.
By far the greater number of the petrified fragments of Calamites afford proof of cambial activity, and possess obvious secondary tissues. In exceptionally perfect specimens the xylem tracheids are found to be succeeded externally by a few flattened thin-walled cells which are in a meristematic condition (fig. 72, A, c); these constitute the cambium zone, and it is the secondary structure that results from the activity of the meristematic cells that we have now to consider.
SECONDARY THICKENING.
In petrified examples of branches in which the secondary thickening has reached a fairly advanced stage, the wood is usually the outermost tissue preserved, the more external tissues having been detached along the line of cambium cells. It is only in a few cases that we are able to examine all the tissues of older examples.