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Invasion Land-Air - Stele

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The term Stele is applied to the tissues that develop from the Procambium.

There has been a progression in the Complexity and LocationStelDiagColr.jpg (109828 bytes) of the Stele throughout the course of Evolution, especially in Stems.

This is largely related to the development of the Megaphyll which has large and numerous Leaf Traces that result in many large Leaf Gaps.

The location of the Stele represents an equilibrium between the location of strengthening and conducting cells.

Engineering studies show that the ideal location for Strengthening Tissue is just beneath the surface of a cylindrical structure (Niklas 1997).

The ideal location for Conducting Tissues is the center of the axis where they would be least likely to break when stems are bent by wind.

It is instructive to note that Roots, which do not produce Leaves, have the most simple stelar types & the stele is in the center of the axis. In most cases this is an Actinostele.

A painful review of the Steles in extant Land Plants

SporophytesPolytrichum240.jpg (47992 bytes)Bryophyta: The "Stele" in the most complex Mosses is located in the center of the stem. It produces minute leaf traces which connect the Stele with the Leaf Vein "Nerve". There are no Leaf Gaps. The Leaves are very small and simple.Strengthening Polytcen240Lab.jpg (57799 bytes)
tissues
occur in the outer Cortex and/or in the "Stele" (Sterids).

BryoStmXsLab.jpg (28572 bytes)
Cross-section through a Moss Stem: Note the thick-walled support tissue in the Cortex. The "Stele" contains thin-walled Hydroids & Leptoids.

Psilophyta: The Stele is located in the center of the stem. EpiphytePR.jpg (58602 bytes)
Leaves lack Vascular Tissues. Leaf Traces are rare or absent. Consequently, there are no Leaf Gaps. The Stele varies from a Haplostele to an Actinostele. Strengthening Tissues are primarily located in the outer Cortex, immediately beneath photosynthetic Parenchyma.

RhizoneX-S240Lab.jpg (52334 bytes)RhizomeStele240Lab.jpg (35583 bytes)

 

 

Psilotum Rhizome with a Haplostele.

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Psilotum Aerial Stem with an Actinostele: Note the Sclerenchyma in the Cortex

Lycophyta: The SteleLycoPerfectDichotPlant.jpg (60485 bytes) is still in the center of the stem. Leaves are Microphylls which contain only one Vein.

Leaf Traces are present in the stem but there are No Leaf Gaps. The Stele varies from a Haplostele to an Actinostele to a Plectostele.Strengthening Tissues are still found in the outer Cortex.

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Haplostele from a Selaginella Root

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Plectostele from a Lycopodium Rhizome

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Protostele from Selaginella: Note the extensive Sclerenchyma development in the Cortex & the numerous Leaf Traces.

Members of the Pterophyta (Ferns) display a wide range of steles. Circin-2Crop.jpg (57644 bytes)
Pterophyta: Thefrillyfern.jpg (102613 bytes) location of the Stele varies from the Center to the Periphery of the Stem. Stele complexity varies from a Haplostele to a Dictyostele to a Polystele.

Megaphylls are present and more than one leaf trace may diverge to each Megaphyll. There can be many overlapping Leafy Gaps which result in a divided stele (Dictyostele). The Vascular Bundles of a Dictyostele are located in the Outer Cortex, rather than in the center of the stem. Rhizomes with no leaves tend to have simple, central Steles (Haplo, Actino, Plecto)

Strengthening Tissue is primarily located in the Outer Cortex.

SteleProtoFernLab400.jpg (66176 bytes)
Haplostele in Fern Rhizome: Note the location of Strengthening Sclerenchyma.
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Fern Siphonostele without Leaf Gaps: Note the location of Strengthening Sclerenchyma.
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Leaf Gaps in the Stele of Dicksonia a Tree Fern
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Dictyostele in Fern Stem
Coniferophyta & Anthophyta: In HerbaceousAbieSnowLolo.jpg (70580 bytes) Plants & the New Growth of Woody Plants, the Stele is found in the Outer Cortex. This is a Eustele.

Monocots have an Atactostele in which most of AnemoneBee.jpg (51116 bytes)the Vascular Bundles are in the outer Cortex but some are present throughout the rest of the stem.

Strengthening Tissue (Sclerenchyma) is associated with the Vascular Bundles rather than the Cortex.

Megaphylls rule and there are many Leaf Gaps in Dicot Steles.

Monocot leaves have many major veins which are directly connected to a complex of highly branched vascular bundles in the stem.

The Bottom Line is that increased Stelar Complexity and its more peripheral Location correlate with the presence of Megaphylls which have large, complex Leaf Traces & many Leaf Gaps in the stem.

Strengthening Tissues are now associated with Vascular Bundles.

The shift from a Eustele to Siphonostele is probably due to a functional shift for each stem segment as new ones are produced by the Shoot Apical Meristem & as Branches get larger.

The Eustele & Atactostele are associated with soft stems that have actively photosynthesizing leaves. These are relatively light-weight and require little structural support.

Furthermore, there is a need for more efficient radial transport out of and into vascular bundles of the leaf. Smaller Vascular Bundles are more efficient in terms of local radial transport than a large Stele located at the center of the stem, or minute Leaf Traces.

The development of a Siphonostele from a Eustele occurs in parts of the stem which have shed their leaves & are far removed from the shoot apex. There is a greater need for structural support in these areas, due to the increased mass that accrues from Primary Growth (new Leaves & Stem tissues). There is also a greater need for Longitudinal Transport to and from the metabolically active parts of the stem tip that is getting further and further away from the Roots.

Structural support is now provided by Secondary Xylem which is located near the periphery of the stem and extends to its center. Most of Xylem conduction occurs in its outer zone ("Sapwood") near the Vascular Cambium & Secondary Phloem.

The more internal "Heartwood" may store excess water for a dry "sunny day".

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Maranta  (Prayer Plant) is Monocot Megaphyll with Net Venation.

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Dicot Stem with a Eustele

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Typical Dicot Vascular Bundle: Note the Sclerenchyma Fibers associated with the Vascular Tissues.


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Cross-section of Makaloa (Cyperus) Stem: This is a Monocot with an Atactostele.


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Vascular Bundles of Makaloa. The blue stain indicates Lignin. Note the great amount of Sclerenchyma which surrounds each bundle!

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Tilia Stem at the end of Primary Growth: There is little Strengthening Tissue present.


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Tilia Stem after one year of growth: Strengthening Tissue (Sclerenchyma) is present in the Secondary Xylem & Secondary Phloem


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Secondary Xylem provides most of  the support for woody plants.

There is a Correlation between the type of Stele and Plant Height. Small Mosses have the most simple Stele. These become more complex and lead to a Eustele for large Herbaceous plants or the growing tips of Woody plants. The Siphonostele develops from a Eustele with Mature Woody stems.

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Roots
typically have Actinosteles with or without Secondary Growth.

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311 Syllabus