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Secondary Phloem


Secondary Phloem has the same origin as secondary xylem, namely, the vascular cambium. Cells displaced towards the outside of the vascular cambium differentiate as phloem. Secondary phloem can remain active over several growth cycles. Secondary phloem, like secondary xylem, is a complex tissue. It always has sieve elements which are analogous to tracheary elements. However, parenchyma is also present, and sclerenchyma may also be visible. There are two kinds of sieve elements, Sieve Cells and Sieve Tube Members.wpe21.jpg (5545 bytes)

Sieve Tube Members are highly specialized for translocation. They are moderately elongate and have horizontal or oblique end walls with sieve plates. Sieve plates contain large sieve pores. The pores are lined with callose which regulates the diameter of the pores. Callose plugs the pores when the cells are ruptured.. The pores can also be closed slowly in response to major changes in the environment, like the onset of winter in the Temperate Zone. Pores closed in this manner may open when growing conditions become favorable. Sieve tubes are formed by the vertical union of several sieve tube members.

Sieve Cells are also specialized for translocation but they lack sieve plates and have a narrower diameter. However, sieve pores are more numerous where sieve cells overlap. Callose is also associated with the sieve pores. Sieve cells are more highly elongated than sieve tube members.

The cell walls of most sieve elements are similar to those of parenchyma cells, and it is often difficult to distinguish between these in cross sections unless a sieve plate or sieve pores can be seen. Callose is preferentially stained by aniline blue. Consequently, sieve elements may be located by their reaction to the stain. However, other cells also react positively with this stain. Callose stained with aniline blue fluoresces under near ultra violet and violet light. This provides a more certain method to locate sieve elements. Callose is also present in their cytoplasm and its fluorescence aids in the location of sieve elements.

Sieve tube members have companion cells, and sieve cells have albuminous cells associated with them. These are also hard to distinguish from phloem parenchyma, especially in cross sections. Starch is usually absent from albuminous cells and this can provide a way to distinguish them.

Rays are present in secondary phloem and are usually continuous with rays in the secondary xylem. Some species develop dilated rays in their secondary phloem. Tilia americana and Hibiscus tiliaceus (hau) are two prime examples for the presence of dilated rays. Dilated rays are principally composed by parenchyma cells which result from localized anticlinal divisions.


Secondary Phloem of Gymnosperms


You should recall that the secondary xylem of gymnosperms like Norfolk Island Pine is simple compared to angiosperms like (Hibiscus tiliaceus). The same is true for secondary phloem.

Important NOTE: Some of the commercial slides have been stained with Lacmoid.

Lignified cell walls will stain blue.

Unlignified cell walls may not stain at all.

Sieve pores, however, stain blue. This accentuates them in secondary phloem.

We will label Lacmoid-stained slides for you!


Secondary Phloem in Gymnosperms



Podocarpus stem cross section at the interface of Secondary Phloem & Xylem stained with Phloroglucinol


Pinus Stem cross-section showing the Epidermis, Cortex, Secondary Phloem & Vascular Cambium.


Pinus Secondary Phloem & Xylem


Secondary Phloem of Pinus


Longitudinal section showing the Secondary Xylem & Phloem of Pinus with typical staining.


Secondary Xylem and Phloem of Pinus stained with Lacmoid

 


Secondary Phloem in Angiosperms


We will use Tilia americana and hau as our principal examples of secondary phloem for angiosperms.

The vascular cambium alternatively produces bands of fibers, phloem parenchyma and sieve tube members plus companion cells.

What is the function of secondary phloem fibers?

Various human civilizations learned to make cloth from bark fibers like those found in Tilia and hau. Hau was not used for the production of kapa by Hawaiians. However they did use the "inner bark" (Secondary Phloem) of Wauke as the source of fibers for kapa. Wauke is also known as Paper Mulberry (Broussonetia papyrifera).

Where is the first phellogen formed?

Where does it develop as the stem enlarges?

Secondary Phloem in Tilia americana


Overview of secondary Phloem in Tilia

 


Cross sections of Tilia stem showing the entire Secondary Phloem, Vascular Cambium & Secondary Xylem




Sieve Tube Member differentiating in Secondary Phloem


Epidermis, Cortex and Secondary Phloem in Tilia. The Periderm has begun to develop! Locate this area!


Same as above but showing the vascular cambium and young secondary xylem.


Find the Companion Cells!

 


Secondary Phloem of Tilia americana viewed with crossed polarizers. What accounts for the birefringence or the lack of birefringence in this image??????

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