ROOTS

10.2: External Root Structure

There are two types of root systems. The first is a fibrous root system which has
multiple big roots that branch and form a dense mass which does not have a visible
primary root (“grass-like”). The other is the taproot system which has one main root
that has branching into lateral roots (“carrot-like”).
Root systems are mainly of two types (Figure 10.2.1). Dicots have a tap root
system, while monocots have a fibrous root system. A tap root system has a main root
that grows down vertically, and from which many smaller lateral roots arise. Dandelions
are a good example; their tap roots usually break off when trying to pull these weeds,
and they can regrow another shoot from the remaining root). A tap root system
penetrates deep into the soil. In contrast, a fibrous root system is located closer to the
soil surface and forms a dense network of roots that also helps prevent soil erosion
(lawn grasses are a good example, as are wheat, rice, and corn). Some plants have a
combination of tap roots and fibrous roots. Plants that grow in dry areas often have
deep root systems, whereas plants growing in areas with abundant water are likely to
have shallower root systems.

(Figure 10.2.1) (a) Tap root systems have a main root that grows down, while (b) fibrous root systems consist of many
small roots. (credit b: modification of work by “Austen Squarepants”/Flickr)

Along with having different systems, there are primary root that originated from
the root of the seedling (Figure 10.2.2𝑎) and secondary (lateral) roots originate from
the primary roots, and adventitious roots originate on stems or leaves, rather than from
the base of the embryo. Adventitious roots can grow if plant cuttings are placed in
water (Figure 10.2.2𝑏).

10.2.2: a) Mung bean tap root (dicot) and wheat bean fibrous root (monocot) from sprouts. b) Jade plant cutting,
showing adventitious roots growing from stem after being placed in water. Credit: Kammy Algiers (CC-BY).

10.3: Root Structure

Root Anatomy
Root growth begins with seed germination. When the plant embryo emerges
from the seed, the radicle of the embryo begins to grow downward and forms the root
system. As the root system grows, various structures begin to appear.
Longitudinal Section
If you were to cut a root down longitudinally, you would see the various layers
inside. The tip of the root is protected by the root cap, a structure exclusive to roots
and unlike any other plant structure. The root cap is continuously replaced because it
gets damaged easily as the root pushes through soil. The root tip can be divided into
three zones: a zone of cell division, a zone of elongation, and a zone of maturation and
differentiation (Figure 10.3.1). The zone of cell division is a continuation of the root
cap; it is made up of the actively dividing cells of the root meristem. The zone of
elongation is where the newly formed cells begin to increase in length, thereby
lengthening the root. They are older than cells in the zone of cell division. Beginning at
the first root hair is the zone of cell maturation where the root cells begin to
differentiate into special cell types. The root has an outer layer of cells called the
epidermis, which surrounds areas of ground tissue and vascular tissue. The epidermis
provides protection and helps in absorption. Root hairs, which are extensions of root
epidermal cells, increase the surface area of the root, greatly contributing to the
absorption of water and minerals. All three zones are in the first centimeter or so of the
root tip.
If you were to cut a cross section of the leaf, you could see other features that are not
as obvious in the longitudinal section. Inside the root, the ground tissue may form two
regions: the cortex and the pith (Figure 10.3.2). When comparing roots to stems, roots
have much more cortex and very little pith. Whereas dicot roots have no central pith,
monocots have a small pith. Both cortex and pith include cells that store photosynthetic
products. The cortex is between the epidermis and the vascular tissue, whereas the
pith lies between the vascular tissue and the center of the root.
The inner portion of the root contains the vascular tissue (xylem and phloem). This
area is called the stele. A layer of cells known as endodermis separates the stele from
the ground tissue in the outer portion of the root (Figure 10.3.2). The endodermis is
exclusive to roots and serves as a checkpoint for materials entering the root’s vascular
system. A waxy substance called suberin is present on the walls of the endodermal
cells. This waxy region, known as the Casparian strips, forces water and solutes to
cross the plasma membranes of endodermal cells instead of slipping between the
cells. This ensures that only materials required by the root pass through the
endodermis, while toxic substances and pathogens are generally excluded. The
outermost cell layer of the root’s vascular tissue is the
pericycle, an area that can give rise to lateral roots.

(Figure 10.3.2): In (left) typical dicots, the vascular tissue forms an X shape in the center of the root. In (right) typical monocots, the
phloem cells and the larger xylem cells form a characteristic ring around the central pith.

Monocots
Note that the size of the stele in the monocot cross section is large (everything inside
the green ring (Figure 10.3.3). The vascular tissue is arranged in a ring around the pith
and are surrounded by the endoderm. The cortext is found outside of the stele. The
exodermis is a layer of cells outside the cortex that controls the flow of water, ions, and
nutrients. The outermost layer of the root is the epidermis, which covers the root and
aids in absorption.
(Figure 10.3.3): Staining reveals different cell types in this light micrograph of a wheat (Triticum) root cross section. Sclerenchyma cells
of the exodermis and xylem cells stain red, and phloem cells stain blue. Other cell types stain black. The stele, or vascular tissue, is the
area inside endodermis (indicated by a green ring). Root hairs are visible outside the epidermis. (credit: scale-bar data from Matt
Russell)
Eudicots
In eudicot roots, the xylem and phloem of the stele are arranged alternately in an X
shape (Figure 10.3.4). Note that there is no pith in a dicot root. Outside the endoderm,
most of the root is composed of cortex tissue. The outer layer is the epidermis.

10.3.4 Dicot root cross section. From center out, the xylem (in red) make an X, and the side tissues (green) make up the phloem..
The endodermis separates the stele from the cortex, which is the largest tissue. The last layer of cells on the edge is the epidermis layer.
(Credit: Wikimedia)

Secondary Root Growth

Many roots have secondary growth as well as primary growth (figures 10.3.5 − 6 ).
This occurs by the production of two types of meristemic tissue, the vascular cambium
and the cork cambium. The cork cambium is responsible for the girth or growth in the
diameter of the root. This occurs by the cork cambium adding vascular tissue to the
root. Cells of the pericycle and procambium (the meristematic tissue between the
primary phloem and xylem) begin division, and form a vascular cambium around the
primary xylem. The vascular cambium then divides to form secondary xylem on the
inside and secondary phloem on the outside.
Figure 10.3.5: A labeled cross section through an older Quercus root, 100x. A=Periderms, B=Seconary Phloem, C=Vascular Cambium, D=Secondary Xylem,
E=Primary xylem. Image from Berkshire Community College Bioscience Image Library (public domain). Labels added by Maria Morrow.

Figure 10.3.6: The process of secondary growth in roots begins when the vascular cambium (dark blue ring) arises from the pericycle and embryonic tissue
called the procamium. The vascular cambium produces secondary xylem (dark red) internally and secondary phloem (light blue) externally. Additionally, the
cork cambium arises from the pericyle and produces cork and phelloderm, forming the periderm (dark brown outer layer). Image from Atlas of Plant and Animal
Histology (CC-BY-NC-SA)

Some roots with secondary growth may form a periderm (a protective layer, replacing
the epidermis). This occurs by the formation of a cork cambium which originate from
the pericycle. The cork cambium produces parenchyma tissues called phelloderm to
the inside of the root and the cork on the outside of the root. Cork cells (phellem) are
dead at maturity. They are hollow and the addition of air space in the tissue functions
as a protective layer. They also produce a waxy substance called suberin. This wax
functions to aid in water loss.
It also makes the root more resistant to bacterial and fungal infections. The three
layers 1. phelloderm 2. cork cambium and 3. cork cells are collectively known as the
periderm.
STEM
11.1 Stem Morphology (External Structure)
Plant stems, whether above- or belowground, are characterized by the
presence of nodes and internodes (figure ). Nodes are points of attachment for
leaves. Leaves often consist of a thin region that attaches to the stem (the
petiole) and a broader blade (see Leaves). The stem region between two
nodes is called an internode. An axillary bud is usually found in the axil— the
area between the base of a leaf and the stem—where it can give rise to a
branch called an axillary shoot (figure ). The shoot apex at the tip shoot
contains the shoot apical meristem surrounded by developing leaves called
leaf primordia (see Meristems).

Figure : Leaves are attached to the plant stem at areas called nodes. An internode is the stem region between two nodes. The smaller leaves just above the nodes are the beginning of axillary
shoots, which arose from axillary buds. Image modified from OpenStax (CC-BY)
Figure : Axillary buds of sweet granadilla (Passiflora ligularis). One of the axillary buds is labeled with an arrow.
Image modified from Eiku (CC-BY-SA).
11.2 Internal Anatomy of the Primary Stem

The primary stem refers to the herbaceous (non-woody) stem, which has not
undergone secondary growth, which produces bark and wood. Some species
(all monocots and some eudicots) remain herbaceous for their entire lives,
maintaining the primary stem. Other species of eudicot initially form a primary
stem but later become woody, replacing the primary stem with the secondary
stem. The anatomy of the stem (internal structure) can be examined through
longitudinal sections (cutting the stem lengthwise) or in cross sections (cutting
a slice of the stem perpendicular to the length).

All three tissue types are represented in the primary stem. The epidermis
is the dermal tissue that surrounds and protects the stem. The epidermis
typically consists of one layer of cells. A waxy cuticle on the outside of these
cells limits water loss. Epidermal cells are the most numerous and least
differentiated of the cells in the epidermis. Pores in the epidermis called
stomata (singular: stoma) allow for gas exchange. Each stoma is bordered by
a pair of guard cells, which regulate stomatal opening. While stomata are
present in stems, they occur at higher densities in leaves. Trichomes are hair-
like structures on the epidermal surface. They help to reduce transpiration (the
loss of water by aboveground plant parts), increase solar reflectance, and store
compounds that defend the leaves against predation by herbivores (figure ).

Figure : Trichomes extend from the epidermis of this Datrua inoxia stem. Image by Silk666 (CC-BY-SA).

Ground tissue fills much of the stem, forming the cortex directly within the
epidermis and the pith (if present) in the center. The outermost portion of the
cortex is usually a few layers of collenchyma cells. The remainder of the cortex
and pith consist of parenchyma cells. The arrangement of vascular tissue in the
stem depends on the species (see below).
Cross sections reveal three possible arrangements of vascular tissue (steles)
in the stem. The first arrangement (solenostele) is present in a few eudicots,
such as basswood (Tilia). In the solenostele, the vascular tissue appears as a
continuous ring (vascular cylinder; figure ). The interfascicular regions (pith
rays) of parenchyma cells that separate vascular tissue are thus extremely
narrow. The second arrangement (eustele) is present in most eudicots such as
sunflower (Helianthus) and buttercup (Ranunculus). In the eustele, vascular
tissue is clustered into distinct vascular bundles arranged in a ring, allowing
for thicker interfascicular regions in between them (figure ). In these
solenosteles and eusteles, the vascular tissue separates ground tissue into an
outer cortex and central pith. The third arrangement (ataktostele) is present in
most monocots, such as corn (Zea mays) and a few eudicots. In the
ataktostele, vascular bundles are scattered throughout the stem (figure ). While
vascular bundles near the outside of the stem are packed more densely in this
third arrangement, their distribution is somewhat disorderly. There is no
distinction between the cortex and pith in the third arrangement.

Figure : Cross section of a common St. John's Wort (Hypericum perforatum) stem, illustrating a silenostele. The central pith (greenish-blue, in the center) and peripheral cortex (narrow
zone 3–5 cells thick just inside the epidermis) are composed of parenchyma cells. Vascular tissue composed of xylem (red) and phloem tissue (green, between the xylem and cortex) surrounds
the pith. Image by RolfDieterMueller (CC-BY)

Figure : In eusteles (left), vascular bundles are arranged around the periphery of the ground tissue. The xylem tissue is located toward the interior of the vascular bundle, and phloem is located
toward the exterior. Primary phloem fibers cap the vascular bundles. In ataktosteles (right), vascular bundles composed of xylem and phloem tissues are scattered throughout the
ground tissue.
Figure : Cross section of a sunflower (Helianthus) stem, illustrating a eustele. Vascular bundles are arranged in a ring.The interfascicular regions between the
vascular bundles are thick compared to the silenostele, where they were too thin to be visible. The ground tissue is separated into an outer cortex and a central
pith. Image labeled from Berkshire Community College Bioscience Image Library (public domain)

Figure : Cross section of a corn (Zea mays) stem, an example of an ataktostele, at 40X magnification. Vascular bundles are scattered but are at a higher density
near the outside of the stem. In contrast to silenosteles and eusteles, the ground tissue is not separated into a pith and cortex. Image labeled from Berkshire
Community College Bioscience Image Library (public domain)

The cells of embryonic tissue called the procambium (see Meristems) divides
to produce primary xylem internally and primary phloem externally. In some
vascular bundles, some procambial cells remain and form the fascicular
cambium in the center of the vascular bundle. Once the stem has finished
lengthening, sclerenchyma fibers called primary phloem fibers are produced
just outside of the primary phloem. The primary phloem fibers of each vascular
bundle are sometimes called phloem caps (bundle caps). If primary phloem
fibers surrounded the entire vascular bundle, they form a bundle sheath (figure
).

Figure : A vascular bundle in the stem of corn (Zea mays) at 400X magnification. The protoxylem vessel element matured prior to the metaxylem vessel elements of the xylem. At maturity,
the protoxylem vessel breaks, leaving an air space. The sieve-tube elements and companion cells form the phloem. The bundle sheath consists of primary phloem fibers (sclerenchyma
fibers) that surround the vascular bundle. Image labeled from Berkshire Community College Bioscience Image Library (public domain)

Vascular bundles connect leaves and stems. The strands of vascular tissue
that connect the leaves to the stem are called leaf traces. Just above leaf
traces are portions of stem without vascular tissue called leaf trace gaps.
Similarly, branch traces connect axillary shoots to the main stem, leaving
branch trace gaps just above them (figure ).

Figure : The purple cylinder represents the vascular tissue in a stem. Vascular tissue leaves the stem to enter the leaf, forming a leaf trace. The space without vascular tissue just above the
leaf trace is the leaf trace gap. The branch trace likewise produces a branch trace gap. Image by Melissa Ha (CC-BY).
Figure : A longitudinal section of a Coleus shoot tip. The leaf trace gaps are spaces without vascular tissue just above the leaf traces, which are strands of vascular tissue moving
from the stem to the leaf. The three tissue system in this stem are represented by the epidermis (outer layer of dermal tissue, not labeled), pith and cortex (ground tissue), and vascular tissue.
Image by Melissa Ha (CC-BY).