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Stems-Primary Growth

Morphology & Anatomy - Monocots

Apical MeristemsAgaveStem80.jpg (8318 bytes)

In many cases there are no differences between Monocot and Dicot Shoot Apical Meristems (SAMs). However some monocots like Coconut Palm, Banana and Ti (Cordyline) have a Meristem which subtends the SAM. Its meristematic cells divide anticlinally near the SAM, and periclinally as they are displaced from the SAM. This  meristem  produce files of cells towards the inside of the stem. This causes an increase in the diameter of the stem in the vicinity of the SAM with virtually no Internodal Elongation. This is called the Primary Thickening Meristem (PTM)and is only found in monocots. Procambium is abundant in the derivative cells of the PTM. These are the primary vascular bundles of the stem. These plants have many veins in their leaves and this is reflected in the number of vascular bundles in the stem.

Primary Thickening Meristem in Ti
CordlineSAM240Lab.jpg (72005 bytes)

BananaSAMHue500.jpg (110442 bytes)

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Primary Thickening Meristem (PTM) in Banana

BananaSAMPrimThickMeiCrop.jpg (24358 bytes)

BananaSAMPrimThickMeiLabb.jpg (69803 bytes)

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Primary Thickening Meristems in Monocots

Monocots like Sugarcane and Corn do not have primary thickening meristems but they do have numerous Vascular Bundles in their leaves and consequently in their stems. The distribution of the Vascular Bundles in monocot stems looks disorganized at first glance. However, they are very well organized, but their organization is complex compared to Dicots.

Typical Monocot Stems

While Asparagus is not the plant usually presented in textbooks, it shows traits which are similar to those in Corn & Sugarcane.

AsparagasXSHueGreen500.jpg (43946 bytes)      AsparagusLab500.jpg (134552 bytes)

Cross-Section of Asparagus Stem - Note the number and size diversity of the Vascular Bundles.

The stem is covered by an Epidermis which has a thick Cuticle & Stomata.

The outer part of the stem is photosynthetic and contains Chlorenchyma and small Vascular Bundles.

This is followed by a zone of thick-walled Sclerenchyma cells.

The rest of the stem is composed of Parenchyma in which Vascular Bundles are embedded.

There is no Pith as vascular bundles can be found near the center of the stem.

There is a progression in the size of the Vascular Bundles from small to large as one proceeds from the outside to the center of the stem.

AsparagusStemPart300.jpg (52961 bytes)

AspStemPartLab400.jpg (59092 bytes)

Outer Part of Asparagus Stem

The Vascular Bundles in Asparagus are a little unusual in that the Xylem almost surrounds the Phloem completely. The Phloem has Exarch development while the xylem is Endarch.

AsparagusStemLgVB.jpg (27921 bytes)

AspVBLab.jpg (35891 bytes)

Large Vascular Bundles in Asparagus

The vascular bundles (VBs) in corn (Zea mays) are "classical" Monocot VBs. They are surrounded by a Bundle Sheath which can be sclerotic. The Phloem has a highly organized appearance. The cells with the greatest diameter are the Sieve Tube Members. The smaller cells are Companion Cells.

The Xylem consists of large Metaxylem Vessel Members and smaller Protoxylem. Corn stems elongate rapidly and many of the protoxylem tracheary elements are   destroyed.  Consequently, the TEs may be missing and a protoxylem cavity (lacuna) can be seen instead.

ZeaVBs.jpg (38662 bytes)

ZeaVBsLab300.jpg (48665 bytes)

Vascular Bundles from Corn Stem - The Classic Monocot

SugCanVBTolBl300.jpg (31320 bytes)

Sugarcane Vascular Bundle Stained with Toluidine Blue Compare with the VB from Corn & Identify the same structures that are labeled above.

Stem of Makaloa (Cyperus laevigatus)

BMHaleClose.jpg (35681 bytes)
Pawehe design for house (Hale) in Makaloa Mat at the Bishop Museum

Ancient Hawaiians used the stems of makaloa to make fine sleeping mats. These are considered to be the finest mats made by Polynesians. Some of these contained Pawehe designs. Mats with Pawehe were made only for royalty. The plants used to make the Pawehe designs were called kohekohe (Eleocharis calva). Both plants belong to the Cyperaceae (Sedge Family), and they grow in semiaquatic environments. We will study the stem anatomy of makaloa. Compare this stem with Asparagus.

The Stem is nearly circular in outline. Photosynthetic tissue is obvious in the outer region of the stem. Vascular bundles are visible at low magnification and they are distributed throughout the stem. There are also prominent air chambers throughout the stem.

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Cross-Section of Makaloa Stem  Unstained

The presence of vascular bundles amid the photosynthetic parenchyma becomes obvious at higher magnification.

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Cross-Section of Makaloa Stem
Unstained

The presence of fiber bundles just beneath the Epidermis becomes apparent at higher magnification.

MakOutStmNoStain400.jpg (15007 bytes)

Outer part of Makaloa Stem

A thick cuticle is observed at higher magnification. The Chlorenchyma cells are distributed around the small, peripheral vascular bundles.

MakEpiFbVbMM400.jpg (15121 bytes)                   MakEpiFbVbMMTolBl300.jpg (21585 bytes)

Epidermis and outer region of Makaloa Stem, Unstained and stained with Toluidine Blue - Note the Cuticle, Fiber Bundles, Chlorenchyma & Small Circular Vascular Bundles.

Staining with Toluidine Blue shows that most of the cell walls are lignified. It also makes the air cavities more obvious, see below.

MakStmXSTolBl400.jpg (27439 bytes)

Makaloa Stem Cross-Section stained with Toluidine Blue  Note the Aerenchyma.

MakLgVBTolBl300.jpg (27980 bytes)

Interior Vascular Bundles of Makaloa Stained with Toluidine Blue Note the Fibers which surround each VB as well as the location of Phloem & Xylem.

Larger Vascular Bundles, similar to those in corn and sugarcane are observed towards the center of the stem. They are surrounded by Bundle Sheaths composed of thick, lignified Sclerenchyma.

Wheat Stem

Wheat is one of the most important crops in the world. Its cultivation is linked to the beginnings of civilization in the Middle-East which is sometimes called the cradle of civilization. The An Eagle is at the center of the Great Seal of the United States of America. He holds shafts of arrows in one talon & shafts of wheat in the other! Wheat cultivation played an important role in the development of the USA across the Great Plains.

WheatStmOuterPart400.jpg (38930 bytes)      WheatStmOuterPartVBs.jpg (43018 bytes)

Cross-Sections of Wheat Stem Note that there are two size classes of Vascular Bundles. Also note the Photosynthetic Parenchyma.

The center part of a mature wheat stem is hollow due to the rapid rate of stem elongation which occurs in stem development. The rim of tissue that remains contains vascular bundles of various sizes. The smaller bundles are associated with chlorenchyma in the outer stem. Larger bundles are seen towards the inside of the stem. Most of the cells may be sclerotic and stain positively for lignin. Stomata are present in the Epidermis. These are associated with the pockets of photosynthetic parenchyma.

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