xylem and protophloem alone are differentiated) being very much shorter than in the stem.
the xylem being centrifugal and the m, Medullary bundles.
Differentiation of the xylem progresses outwards, of the phloem inwards, but the two tissues never meet in the centre.
The vascular bundles of the stem belong to the col xylem and the bast or phloem stand side by side on the same radius.
occasionally happens that groups of (After Worsdell.) xylem and phloem are developed pd, Periderm in leaf-bases.
the xylem coming to the surface of the cylinder, ~dS;
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.
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).
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.
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 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.
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.
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.
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 tracheids or vessels, indifferently called tracheal elements, together with the immediately associated cells (usually amylom in Pteridophytes) constitute the xylem of the plant.
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.
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 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 latter is often sclerized, especially opposite the phloem, and to a less extent opposite the xylem, as in the stem.
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.
The vascular system of cycadean seedlings presents some features worthy of note; centripetal xylem occurs in the cotyledonary bundles associated with transfusion-tracheids.
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.
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.
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.
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.
(After the xylem recalls the Fujii.) cycadean type.
Such a vascular cylinder is called a haplostele, and the axis containing it is said to be haplostelic. In the stele of the root the strands of tracheids along the lines where the xylem touches the pericycle are spiral or annular, and are the xylem elements first formed when the cylinder is developing.
The xylem and phloem also, rarely form perfectly continuous layers as they do in a solenostelic fern.
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.
The cylinder is surrounded by a mantle of one or more layers of parenchymatous cells, the pericycle, and the xylem is generally separated from the phloem in the stem by a similar layer, the mesocycle (corresponding with the amylom sheath in mosses).
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.
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.
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 parenchyma is often found in strands associated with the tracheal elements.
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.
intercellular channel representing xylem; ph.
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.
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.
metu starchy xylem-parenchyma, which, when the xylem is bulky, usually appear among the tracheids, the phloem also often being penetrated by similar bands of phloem-parenchyma.)
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.
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.
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.
In each bundle, separating the xylem and phloem, is a layer of meristem or active formative tissue, known as cambium; by the formation of a layer of cambium between the bundles (interfascicular cambium) a complete ring is formed, and a regular periodical increase in thickness results from it by the development of xylem on the inside and phloem on the outside.
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.
Additionally, xylem development, or xylogenesis, has a significant impact on the value of many crop species.
the leptom found in Pteridophytes and Phanerogams with its associated cells, and is entirely parallel with the xylem.
As the primitive stele of a Pteridophyte is traced upwards from the primary rout into the stem, the phloem becomes continuous round the xylem.
In others a central parenchyma or primetive pith a new region of the primitive stelar conjunctiveappears in the centre of the xylem.
All the metaxylems join at the nodes into a complete ring of xylem.
Sometimes in such cases the cambium ceases to be active round these bays and joins across the outside of the bay, where it resumes its normal activity, thus isolating a phtoem strand, or, as it is sometimes called, a phloem -island, in the midst of the xylem.
An ordinary cambium is scarcely ever found in the Monocotyledons, but in certain woody forms a secondary meristem is formed outside the primary bundles, and gives rise externally to a little secondary cortex, and internally to a secondary parenchyma in which are developed numerous zones of additional bundles, usually of concentric structure, with phloem surrounded by xylem.
Cases of complete fusion occur in the formation of laticiferous vessels, and in the spiral, annular and reticulate vessels of the xylem.
inverse orientation of the tissues, x, Xylem.
C, seminiferous scale as fused pair of leaves (1 1, 1 2, l 3, first, second and third leaves; b, shoot; Br, bract), D, cone-scale of Araucaria (n, nucellus; i, integument; x, xylem).
When tracheids occur in the medullary rays of the xylem these are replaced in the phloem-region by irregular parenchymatous cells known as albuminous cells.
Resin-canals, which occur abundantly in the xylem, phloem or cortex, are not found in the wood of the yew.
It has been suggested that transfusion-tracheids represent, in part at least, the centripetal xylem, which forms a distinctive feature of cycadean leaf-bundles; these short tracheids form conspicuous groups laterally attached to the veins in Cunninghamia, abundantly represented in a similar position in the leaves of Sequoia, and scattered through the so-called pericycle in Pinus, Picea, &c. It is of interest to note the occurrence of precisely similar elements in the mesophyll of Lepidodendron leaves.
The vascular bundles themselves are collateral, the xylem consisting of the protoxylem, towards the centre of the stem, and two groups of xylem, between which the phloem is situated; the protoxylem elements soon break down, giving rise to the carinal canal.
Only the median or carinal strand of xylem is common to stem and leaf; the lateral cauline strands possibly represent the remains of a centripetally developed mass of primary xylem.
The central cylinder of the root, in which there are several xylem and phloem strands, has around it a two-layered endodermis, the inner layer of which appears to take the place of a pericycle.
The single stele in the stem consisted of the phloem surrounding a solid central strand of xylem, the groups of protoxylem being situated at the projecting angles.
In Sphenophyllum, in which the transverse section of the xylem is triangular, there were three or six protoxylem groups; in Cheirostrobus they were more numerous.
The stem is monostelic, the protoxylem groups being towards the periphery of the xylem, the development of which is thus centripetal; the centre of the stele is occupied by sclerenchymatous tissue.
The anatomy of Lycopodium presents considerable variety in detail, but the stem is always monostelic and the development of the xylem centripetal, the protoxylems being situated at the periphery of the stele; pericycle and endodermis surround the stele, and the wide cortex may be more or less sclerenchymatous.
The phloem completely surrounds the xylem, which usually develops from two protroxylem groups.
The stem had a single stele, the primary xylem of which was polyarch and centripetally developed.
A meristematic zone forms a short distance outside the xylem, from which secondary tissue is developed both internally and externally; that to the inside contains both xylem and phloem elements.
The stem is monostelic, the arrangement of the xylem and phloem being collateral.
In the roots of Ophioglossum and Botrychium and in the first formed roots of Helminthostachys an endophytic fungus is present, forming a mycorhiza - the stele in the larger roots has the usual radial arrangement of xylem and phloem; monarch roots occur in Ophioglossum.
The stem, from the ground tissue of which sclerenchyma is absent, has a complicated system of steles arranged in concentric circles; the thick roots, the central cylinders of which have several alternating groups of xylem and phloem, arise in relation to these.
The structure is in all respects that typical of roots, as shown by the centripetal primary wood, and the alternation of xylem and phloem groups observable in exceptionally favourable young specimens.
The primary structure f ` is only found unaltered in the 1 1, 1 ` by means of a cambium set in very Il 111/1/1, 1 1 early, xylem being formed internally and phloem externally in a perfectly normal manner.
The stem is traversed by a single stele, with solid wood, without pith; the primary xylem is triangular in section, the spiral elements forming one or two groups at each angle, while the phloem occupied the bays, so that the structure resembles that of a triarch root.
In other species, however, the centripetal primary xylem is represented.
The leaf-traces, where they traverse the cortex, have the structure of the foliar bundles in Cycads, for they are of the collateral type, and their xylem is mesarch, the spiral elements lying in the interior of the ligneous strand.
Corresponding strands of primary xylem have been observed in stems of the genus Pitys (Witham), of Lower Carboniferous age, which consisted of large trees, probably closely allied to Cordaites.
More specifically, our results point to the involvement of these MYB proteins in the regulation of the formation of lignin during xylem formation.
We have cloned two cDNAs which encode the only members of the Myb class of transcription factors found in differentiating pine xylem.
Movement in xylem vessels The cells which make up a xylem vessels The cells which make up a xylem vessel are dead.
xylem formation in pine by cDNA sequencing.
xylem tissue are usually strengthened with lignin, the woody material we are familiar with in trees and shrubs.
xylem cells and across the leaf.
xylem development, or xylogenesis, has a significant impact on the value of many crop species.
In the primary axis of the plant among Pteridophytes and many Phanerogams, at any rate in its first formed part, the xylem and phloem are associated in the form of a cylinder (stele), with xylem occupying the centre, and the phloem (in the upward-growing part or primary stem) forming a mantle at the periphery (fig.
In the root the mesocycle, like the phloem, is interrupted, and runs into the pericycle where the xylem touches the latter (fig.
The stele of most Lycopods is amore or less modified protostele, but in the genus Lyco podium a peculiar arrangement of the xylem Ab and phloem is found, in which the latter, instead of being erran confined to a peripheral mantle of tissue, forms bands Stelar running across the stele and alternating with similal Systems of bands of xylem (fig.
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.
While the stele of the primary root in both Gymnosperms and Angiosperms is usually diarch or tetrarch, the large primary root-steles of many adventitious roots are frequently polyarch, sometimes with a very large number of protoxylems. Such a stale seldom has the centre filled up with xylem, this being replaced by a large-celled pith, so that a siphonostelic structure is acquired (fig.
The camb-ium in the typical case, which is by far the most frequent, continues the primary differentiation of xylem and phloem in the desmogen strand (see above), or arises in the resting mesodesm or mesocycle and adds new (secondary) xylem and phloem to the primary tissues.
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.
The xylem parenchyma cells are connected, as are the medullary ray cells, with the tracheal elements by one-sided bordered pitsi.e.
The top and bottom rows of the xylem rays are often developed as irregularly-thickened radially-elongated tracheids which serve for the radial conduction of water, and communicate with the ordinary tracheids of the secondary xylem by large bordered pits.
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.
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.
These concentric rings of secondary xylem and phloem (fig.
A peculiarity of these leaves is the inverse orientation of the vascular tissue; each of the two veins has its phloem next the upper and the xylem towards the lower surface of the leaf; this unusual position of the xylem and phloem may be explained by regarding the needle of Sciadopitys as being composed of a pair of leaves borne on a short axillary shoot and fused by their margins (fig.
The rootlets, which branched by dichotomy, contain a slender monarch stele exactly like that in the roots of Isoetes and some Selaginellae at the present day; they possessed, however, a complex absorptive apparatus, consisting of lateral strands of xylem, connecting the stele with tracheal plates in the outer cortex.
Movement in xylem vessels The cells which make up a xylem vessel are dead.
Analysis of xylem formation in pine by cDNA sequencing.
The hollow tubes of xylem tissue are usually strengthened with lignin, the woody material we are familiar with in trees and shrubs.
This is turn sets up a concentration gradient across which water moves by osmosis out of the xylem cells and across the leaf.
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