C/i, epidermis; st stoma; me,, mesophyil; pal, palisade; spa, spongy tissue; Isp, inteicellular space; wi., water tissue; x, xylem; p/i, phioem; Phil, phloeoterma; sri, scierenchyma.
The xylem and phloem are nearly always found in close association in strands of various shapes in all the three main organs of the sporophyteroot, stem and leafand form a connected tissue-system running through the whole body.
The xylem coming to the surface of the cylinder, ~dS;
Each strand of spiral or annular first-formed tracheids is called a protoxylem strand, as distinct from the metaxylem or rest of the xylem, which consists of thick-walled tracheids, the pits of which are often scalariform.
When there is a single protoxylem strand in the centre of the stele, or when, as is more commonly the case, there are several protoxylem strands situated at the internal limit of the xylem,, the centre of the stem being occupied by parenchyma, the stele is endarch.
This consists of a few xylem elements, e a a segment of phloem, pericycle, and usually an arc of h~s endodermis, which closes round the bundle as it detaches ~
Where internal phloem is present this is separated from the internal endodermis by an endocycle or internal pericycle, as it is sometimes called, and from the xylem by an internal mesocyclethese two layers, together with the outer mesocycle and pericycle, constituting the conjunctive tissue of the now hollow cylindrical stele.
It is probable that this type of stele is a modification of a primitive protostele, in which the main mass of stelar xylem has become much reduced and incidentally separated from the leaftraces.
The main events in this transition appear to have been (I) disappearance of the central xylem of the protostele and replacement by pith, leading to the survival of a number of (mesarch) collateral bundles (see below) at the periphery of the stele; (2) passage from mesarchy to endarchy of these bundles correlated with a great increase in secondary thickening of the stele.
The xylem and phloem also, rarely form perfectly continuous layers as they do in a solenostelic fern.
13, 23), the xylem of which is usually wedgeshaped in cross-section with the protoxylem elements at the inner extremity, while the phloem forms a band on the outer side of the xylem, and separated from it by a band of conjunctive tissue (mesodesm).
In the stems of many water-plants various stages of reduction of the vascular system, especially of the xylem, are met with, and very often this reduction leads to the formation of a compact stele in which the individuality of the separate Reduced bundles may be suppressed, so that a closed cylinder lmpbost~h1c of xylem surrounds a pith.
The leaf-bundles are always collateral (the phloem being turned downwards and the xylem upwards), even in Ferns, where the petiolar strands are concentric, and they have the ordinary mesodesm and peridesm of the collateral bundle.
The latter is often sclerized, especially opposite the phloem, and to a less extent opposite the xylem, as in the stem.
Differentiation of the xylem progresses outwards, of the phloem inwards, but the two tissues never meet in the centre.
Xylem and protophloem alone are differentiated) being very much shorter than in the stem.
New tangential walls arise in the cells which are the seat of cambial activity, and an initial layer of cells is established which cuts off tissue mother-cells on the inside and outside, alternately contributing to the xylem and to the phloem.
A tissue mother-cell of the xylem may, in the most advanced types of Dicotyledons, give rise to(I) a tracheid; (2) a segment of a vessel; (3) a xylem-fibre; or (4) a vertical file of xylem-parenchyma cells.
At celtain points the cambium does not give rise to xylem and phloem elements, but cuts off cells on both sides which elongate radially and divide by horizontal walls.
When a given initial cell of the cambium has once begun to produce cells of this sort it continues the process, so that a radial plate of parenchyma cells is formed stretching in one plane through the xylem and phloem.
The xylem and phloem parenchyma consist of living cells, fundamentally similar in most respects to the medullary ray cells, which sometimes replace them altogether.
The xylem parenchyma is often found in strands associated with the tracheal elements.
The xylem parenchyma cells are connected, as are the medullary ray cells, with the tracheal elements by one-sided bordered pitsi.e.
A considerable evolution in complexity can be traced in passing from the simplest forms of xylem and phloem found in the primary vascular tissues both among Pteridophytes and Phanerogams to these highly differentiated types.
Among Gymnosperms the secondary xylem is similarly simple, consisting of tracheids which act as stereom as well as hydrom, and a little amylom; while the phloem-parenchyma sometimes undergoes a differentiation, part being developed as amylom, part as proteid cells immediately associated with the sieve-tube, in other cases the proteid cells of the secondary phloem do not form part of the phloem-parenchyma, but occupy the top and bottom cellrows of the medullary rays, the middle rows consisting of ordinary starchy cells.
Wood thus altered is known as heart-wood, or duramen, as distinguished from the young sap-wood, or alburnum, which, forming a cylinder next the cambium, remains alive and carries on the active functions of the xylem, particularly the conduction of water.
Where the primary bundles are farther apart, so that the primary rays are wider, the interfascicular cambium may form several fairly broad (principal) secondary rays in continuation of certain radial bands of the primary ray, and between these, wedges of secondary xylem and phloem: or, finally, secondary xylem and phloem may be formed by the whole circumference ot the cambium, fascicular and interfascicular alike, interrupted only by narrow secondary rays, which have no relation to the primary ones.
In the simplest cases the cambium produces xylem more freely along certain tracts of the circumference than along others, so that the stem loses its original cylindrical form and becomes elliptical or lobed in section.
In others the secondary phloem is produced more abundantly in those places where the secondary xylem is deficient, so that the stem remains cylindrical in section, the phloem occupying the bays left in the xylem mass.
The formation of additional cambial cylinders or bands occurs in the most various families of Dicotyledons and in some Gymnosperms. They may arise in the pericycle or endocycle of the stele, in the cortex of the stem, or in the parenchyma of the secondary xylem or phloem.
Sometimes the activity of the successive cambiums simply results in the formation of concentric rings or arcs of secondary xylem and phloem.
The cambium in the root, which is found generally in those plants which possess a cambium in the stem, always begins in the conjunctive tissue internal to the primary phloems, and Camblum forms new (secondary) phloem in contact with the In Roots primary, and secondary xylem internally.
If the development of secondary tissues is to proceed further, arcs of cambium are formed in the pericycle external to the primary xylems, and the two sets of cambial arcs join, forming a conti,riuous, wavy line on transverse section, with bays opposite the primary phloems and promontories opposite the primary xylems. Owing to the resistance offered by the hard first-formed secondary xylem, the bays are pushed outwards as growth proceeds, and the wavy line becomes a circle.
Opposite the primary xylems, the cambium either (a) forms parenchyma on both sides, making a broad, secondary (principal) ray, which interrupts the vascular ring and is divided at its inner extremity by the islet of primary xylem; or (b) forms secondary xylem and phloem in the ordinary way, completing the vascular ring.
The vascular bundles of the stem belong to the col xylem and the bast or phloem stand side by side on the same radius.
Secondary xylem and phloem produced by a single cambium, or by successive cambial zones; no true vessels (except in the Gnetales) in the wood, and no companioncells in the phloem.
After the cambium has been active for some time producing secondary xylem and phloem, the latter consisting of sievetubes, phloem-parenchyma and frequently thick-walled fibres, a second cambium is developed in the pericycle; this produces a second vascular zone, which is in turn followed by a third cambium, and so on, until several hollow cylinders are developed.
Occasionally happens that groups of (After Worsdell.) xylem and phloem are developed pd, Periderm in leaf-bases.
The xylem being centrifugal and the m, Medullary bundles.
Most of these cortical bundles are collateral in structure, but in some the xylem and phloem are concentrically arranged; the secondary origin of these bundles from procambium-strands was described by Mettenius in his classical paper of 1860.
A leaf-trace, as it passes through the cortex, has a collateral structure, the protoxylem being situated at the inner edge of the xylem; when it reaches the leaf-base the position of the spiral tracheids is gradually altered, and the endarch arrangement (protoxylem internal) gives place to a mesarch structure (protoxylem more or less central and not on the edge of the xylem strand).
In a bundle examined in the basal portion of a leaf the bulk of the xylem is found to be centrifugal in position, but internally to the protoxylem there is a group of centripetal tracheids; higher up in the petiole the xylem is mainly centripetal, the centrifugal wood being represented by a small arc of tracheids external to the protoxylem and separated from it by a few parenchymatous elements.
Finally, in the pinnae of the frond the centrifugal xylem may disappear, the protoxylem being now exarch in position and abutting on the phloem.
The vascular system of cycadean seedlings presents some features worthy of note; centripetal xylem occurs in the cotyledonary bundles associated with transfusion-tracheids.
(After the xylem recalls the Fujii.) cycadean type.
A point of anatomical interest is the occurrence in the vascular bundles of the cotyledons, scale-leaves, and elsewhere of a few centripetally developed tracheids, which give to the xylem-strands a mesarch structure such as characterizes the foliar bundles of cycads.
In 1869 van Tieghem laid stress on anatomical evidence as a key to the morphology of the cone-scales; he drew attention to the fact that the collateral vascular bundles of the seminiferous scale are inversely orientated as compared with those of the carpellary scale; in the latter the xylem of each bundle is next the upper surface, while in the seminiferous scale the phloem occupies that position.
The roots of many conifers possess a narrow band of primary xylem-tracheids with a group of narrow spiral protoxylem-elements at each end (diarch).
The annual rings in a root are often less clearly marked than in the stem, and the xylem-elements are frequently larger and thinner.
It is in the nature of the secondary xylem that the Coniferales are most readily distinguished from the Dicotyledons and Cycadaceae; the wood is homogeneous in structure, consisting almost entirely of tracheids with circular or polygonal bordered pits on the radial walls, more particularly in the late summer wood.