Handbook of Plant Science root covered by cuticle but, there is a different muatan dari wax-nya

shg msh bs permeable

Plants can grow indeterminately. The ongoing growth of roots is made possible by the capability of
the plant to maintain a pool of stem cells. These stem cells keep proliferating and do not differentiate.
The daughters of the stem cells differentiate and form the different structures of the root. In the plant
root, stem cells are located in the root tip. The stem cells are located in an organized structure, the
stem cell niche. The stem cell niche functions as a microenvironment where, by short range cell
signaling, the stem cells are maintained. The Quiescent Center (QC) lies in the centre of the root stem
cell niche and provides the signals for maintenance of the surrounding stem cells. The organization of
the niche is set up early in embryogenesis and stays unchanged in the mature plant. Essential for root
patterning in embryogenesis is the phytohormone auxin. Several transcription factors are involved in
both root patterning in the embryo and stem cell maintenance in the mature plant. The processes of
organization and maintenance of root stem cells are very well studied in the Arabidopsis thaliana and
new insights from this model organism will be discussed here.



The root system of a plant constantly provides the stems and leaves with water and dissolved
minerals. In order to accomplish this the roots must grow into new regions of the soil. The
growth and metabolism of the plant root system is supported by the process of photosynthesis
occurring in the leaves. Photosynthate from the leaves is transported via the phloem to the
root system. Root structure aids in this process. This section will review the different kinds
of root systems an look at some specialized roots, as well as describe the anatomy of the roots
in monocots and dicots.
Root Systems:
Taproot System:
Characterized by having one main root (the taproot) from which smaller branch roots
emerge. When a seed germinates, the first root to emerge is the radicle, or primary root. In
conifers and most dicots, this radicle develops into the taproot. Taproots can be modified for
use in storage (usually carbohydrates) such as those found in sugar beet or carrot. Taproots
are also important adaptations for searching for water, as those long taproots found in mesquite
and poison ivy.


Fibrous Root System:
Characterized by having a mass of similarly sized roots. In this case the radicle from a
germinating seed is short lived and is replaced by adventitious roots. Adventitious roots are
roots that form on plant organs other than roots. Most monocots have fibrous root
systems. Some fibrous roots are used as storage; for example sweet potatoes form on fibrous
roots. Plants with fibrous roots systems are excellent for erosion control, because the mass of
roots cling to soil particles.

Root Structures and Their Functions:

Root Tip: the end 1 cm of a root contains young tissues that are divided into the root cap,
quiescent center, and the subapical region.
Root Cap: root tips are covered and protected by the root cap. The root cap cells are derived
from the rootcap meristem that pushes cells forward into the cap region. Root cap cells
differentiate first into columella cells.Columella cells contain amylopasts that are responsible
for gravity detection. These cells can also respond to light and pressure from soil
particles. Once columella cells are pushed to the periphery of the root cap, they differentiate
into peripheral cells. These cells secrete mucigel, a hydrated polysaccharide formed in the
dictyosomes that contains sugars, organic acids, vitamins, enzymes, and amino acids. Mucigel
aids in protection of the root by preventing desiccation. In some plants the mucigel contains
inhibitors that prevent the growth of roots from competing plants. Mucigel also lubricates the
root so that it can easily penetrate the soil. Mucigel also aids in water and nutrient absorption
by increasing soil:root contact. Mucigel can act as a chelator, freeing up ions to be absorbed by
the root. Nutrients in mucigel can aid in the establishment of mycorrhizae and symbiotic
bacteria.
Quiescent Center: behind the root cap is the quiescent center, a region of inactive cells. They
function to replace the meristematic cells of the rootcap meristem. The quiescent center is
also important in organizing the patterns of primary growth in the root.
Subapical Region: this region, behind the quiescent center is divided into three zones. Zone of
Cell Division - this is the location of the apical meristem (~0.5 -1.5 mm behind the root
tip). Cells derived from the apical meristem add to the primary growth of the root. Zone of
Cellular Elongation - the cells derived from the apical meristem increase in length in this
region. Elongation occurs through water uptake into the vacuoles. This elongation process
shoves the root tip into the soil. Zone of Cellular Maturation - the cells begin
differentiation. In this region one finds root hairs which function to increase water and
nutrient absorption. In this region the xylem cells are the first of the vascular tissues to
differentiate.

Mature Root: the primary tissues of the root begin to form within or just behind the Zone of
Cellular Maturation in the root tip. The root apical meristem gives rise to three primary
meristems: protoderm, ground meristem, and procambium.


Epidermis: the epidermis is derived from the protoderm and surrounds the young root one cell
layer thick. Epidermal cells are not covered by cuticle so that they can absorb water and
mineral nutrients. As roots mature the epidermis is replaced by the periderm.
Cortex: interior to the epidermis is the cortex which is derived from the ground meristem. The
cortex is divided into three layers: the hypodermis, storage parenchyma cells, and the
endodermis. The hypodermis is the suberinized protective layer of cells just below the
epidermis. The suberin in these cells aids in water retention. Storage parenchyma cells are
thin-walled and often store starch. The endodermis is the innermost layer of the
cortex. Endodermal cells are closely packed and lack intercellular spaces. Their radial and
transverse walls are impregnated with lignin an suberin to form the structure called the
Casparian Strip. The Casparian Strip forces water and dissolved nutrients to pass through the
symplast (living portion of the cell), thus allowing the cell membrane to control absorption by the
root.
Stele: all tissues inside the endodermis compose the stele. The stele includes the outer most
layer, pericycle, and the vascular tissues. The pericycle is a meristematic layer important in
production of branch roots. The vascular tissues are made up of the xylem and phloem. In
dicots the xylem is found as a star shape in the center of the root with the phloem located
between the arms of the xylem star. New xylem and phloem is added by the vascular cambium
located between the xylem and phloem. In monocots the xylem and phloem form in a ring with s
the central portion of the root made up of a parenchymatous pith.
Why does the epidermis on a leaf have no chloroplasts
in it?
The epidermis has numerous functions: protection against various chemical and physical influences,
against being fed upon by animals and against infestation by parasites protection of the plant against
desiccation participation in gas exchange, in secretion of metabolic compounds and in absorption of
water
site of receptors for light and mechanical stimuli that help to transform signals from the surrounding to
the plant
The epidermis has accordingly a number of differentiated cell types to serve the various
functions.Variations typical for certain species and different organizations of the epidermis in the
miscellaneous plant organs add to the number of different cells. Three main types exist:
1. the basic epidermis cell
2. the cells of the stoma complexes and
3. the trichomes (gr.: trichoma = hair), epidermal attachments of varying shape, structure and
function

The basic epidermis cells, i.e. the least specialized cells constitute the largest group of dermal cells.
They seem either polygonal or elongated in top view. Their walls are often wavy or sinuate. It is
unknown, what induces this shape during development, since the explanations given by the existing
hypotheses seem insufficient. Elongated epidermis cells can be found at organs or parts of organs
that are elongated themselves, like stems, leaf petioles, leaf veins or leaves of monocots. The
epidermises of the leaf's upper- and undersurface may have different structures. The shape of the
cells, the thickness of the walls as well as the distribution and number of specialized cells (guard cells
and trichomes) per area may all vary. Wide varieties of different cell shapes may even exist in species
of a single family, e. g. in the Crassulaceae family

The wall of epidermal cells that constitute the leaf's surface is often thicker than the other walls. This
can be particularly well observed with the epidermis of conifer needles and that of xerophytes (plants
living in dry habitats). Aquatic plants have usually thin walls. The wall of many seeds becomes
stronger during ripening and may fill nearly all of the cell's lumen so that the protoplast is driven out
and degenerates. The basic epidermal cells of most species contain no chloroplasts. Some ferns and
several aquatic or shade plants are exceptions.

The epidermis is more often than not built from a single cell layer, though multi-layered, water-storing
epidermises that evolved from initially single-layered tissues by periclinal division have been shown
among the species of several families (Moraceae: most Ficus-species, Piperaceae: Peperonia,
Begoniaceae, Malvaceae and others). Epidermis cells secrete a cuticle, that covers all epidermal
surfaces like an uninterrupted film. It may either be smooth or structured by bulges, rods, filaments,
folds, or furrows.
Although leaves are green most of the epidermal cells do not have functional chloroplasts.
In most plants guard cells are the only cells within a leaf's epidermis that contain chloroplasts
It does not function as a photosynthesizing tissue since it has various other primary functions thus in
the process of evolution to increase its efficiency it may have lost its chloroplast
Although it should be noted that it is not entirely true that epidermal cell do not have chlp there are al
The lack of chloroplast is due to the function of the epidermis. The epidermis is transparent; to allow
light to enter, it's waxy to prevent water loss due to evaporation and it is the first barrier for the leaf
from the external environment very much like human skin. The epidermis protects, regulates and
keeps the leaf airtight.
The absence of chloroplast in the epidermal layers clearly indicate the function of epidermis is not to
prepare food but to protect the internal tissues.

The epidermis cells (from the Greek "", meaning "over-skin") is a single-layered group of
cells that covers plants' leaves, flowers, roots and stems. It forms a boundary between the plant and
the external environment. The epidermis serves several functions, it protects against water loss,
regulates gas exchange, secretes metabolic compounds, and (especially in roots) absorbs water and
mineral nutrients. The epidermis of most leaves shows dorsoventral anatomy: the upper (adaxial) and
lower (abaxial) surfaces have somewhat different construction and may serve different functions.
Woody stems and some other stem structures produce a secondary covering called the periderm that
replaces the epidermis as the protective covering.
Contents
[hide]
1 Description
2 Guard cells
3 Cell differentiation in the epidermis
4 See also
5 References
6 External links
Description[edit]
The epidermis is the outermost cell layer of the primary plant body. In some older works the cells of
the leaf epidermis have been regarded as specialised parenchyma cells,
[1]
but the established modern
preference has long been to classify the epidermis as dermal tissue, whereas parenchyma is
classified as ground tissue.
[2]
The epidermis is main component of the dermal tissue system of leaves
(diagrammed below), and also stems, roots, flowers, fruits, and seeds; it is
usually transparent (epidermal cells have a lower number of chloroplasts or lack them completely,
except for the guard cells.)
The cells of the epidermis are structurally and functionally variable. Most plants have an epidermis
that is a single cell layer thick. Some plants like Ficus elastica and Peperomia, which have periclinal
cellular division within the protoderm of the leaves, have an epidermis with multiple cell layers.
Epidermal cells are tightly linked to each other and provide mechanical strength and protection to the
plant. The walls of the epidermal cells of the above ground parts of plants contain cutin, and are
covered with a cuticle. The cuticle reduces water loss to the atmosphere, it is sometimes covered
with wax in smooth sheets, granules, plates, tubes or filaments. The wax layers give some plants a
whitish or bluish surface color. Surface wax acts as a moisture barrier and protects the plant from
intense sunlight and wind.
[3]
The underside of many leaves have a thinner cuticle than the top side,
and leaves of plants from dry climates often have thickened cuticles to conserve water by reducing
transpiration.

The epidermal tissue includes several differentiated cell types: epidermal cells, guard cells, subsidiary
cells, and epidermal hairs (trichomes). The epidermal cells are the most numerous, largest, and least
specialized. These are typically more elongated in the leaves of monocots than in those of dicots.

Trichomes or hairs grow out from the epidermis in many species. In root epidermis, epidermal hairs,
termed root hairs are common and are specialized for absorption of water and mineral nutrients.
In plants with secondary growth, the epidermis of roots and stems is usually replaced by a periderm
through the action of a cork cambium.
Guard cells[edit]


Stoma in a tomato leaf (microscope image)
Main article: Stoma
The leaf and stem epidermis is covered with pores called stomata (sing., stoma), part of a stoma
complex consisting of a pore surrounded on each side by chloroplast-containing guard cells, and
two to four subsidiary cells that lack chloroplasts. The stoma complex regulates the exchange of
gases and water vapor between the outside air and the interior of the leaf. Typically, the stomata are
more numerous over the abaxial (lower) epidermis of the leaf than the (adaxial) upper epidermis. An
exception is floating leaves where most or all stomata are on the upper surface. Vertical leaves, such
as those of many grasses, often have roughly equal numbers of stomata on both surfaces. The stoma
is bounded by two guard cells. The guard cells differ from the epidermal cells in the following aspects:
The guard cells are bean-shaped in surface view, while the epidermal cells are irregular in shape
The guard cells contain chloroplasts, so they can manufacture food by photosynthesis (The
epidermal cells do not contain chloroplasts)
Guard Cells are the only epidermal cells that can make sugar. According to one theory, in
sunlight the concentration of potassium ions (K+) increases in the guard cells. This, together with
the sugars formed, lowers the water potential in the guard cells. As a result, water from other cells
enter the guard cells by osmosis so they swell and become turgid. Because the guard cells have
a thicker cellulose wall on one side of the cell, i.e. the side around the stomatal pore, the swollen
guard cells become curved and pull the stomata open.
At night, the sugar is used up and water leaves the guard cells, so they become flaccid and the
stomatal pore closes. In this way, they reduce the amount of water vapour escaping from the leaf.
Cell differentiation in the epidermis[edit]


Scanning electron microscope image ofNicotiana alata leaf's epidermis, showingtrichomes (hair-like appendages)
andstomata (eye-shaped slits, visible at full resolution)
The plant epidermis consists of three main cell types: pavement cells, guard cells and their subsidiary
cells that surround the stomata and trichomes, otherwise known as leaf hairs. The epidermis of petals
also form a variation of trichomes called conical cells. These cells all develop from the pavement
cells, which make up the majority of the plants surface cells. In short, cellular differentiation of the
epidermal cells is controlled by two major factors:genetics and environmental conditions.
Trichomes develop at a distinct phase during the actual leaf development, under the control of two
major trichome specification genes: TTG and GL1. The process may be controlled by the plant
hormones gibberellins, and even if not completely controlled, gibberellins certainly have an effect on
the development of the leaf hairs. GL1 causes endoreplication, the replication of DNA without
subsequent cell division as well as cell expansion. GL1 turns on the expression of a second gene for
trichome formation, GL2, which controls the final stages of trichome formation causing the cellular
outgrowth.
Arabidopsis thaliana uses the products of inhibitory genes to control the patterning of trichomes, such
as TTG and TRY. The products of these genes will diffuse into the lateral cells, preventing them from
forming trichomes and in the case of TRY promoting the formation of pavement cells.
As previously mentioned, conical cells are a form of trichome that occurs on the petals of flowers.
Expression of the gene MIXTA, or its analogue in other species, later in the process of cellular
differentiation will cause the formation of conical cells over trichomes. MIXTA is a transcription factor.
Stomatal patterning is a much more controlled process, as the stoma effect the plants water retention
and respiration capabilities. As a consequence of these important functions, differentiation of cells to
form stomata is also subject to environmental conditions to a much greater degree than other
epidermal cell types.
Stomata are holes in the plant epidermis that are surrounded by two guard cells, which control the
opening and closing of the aperture. These guard cells are in turn surrounded by subsidiary
cells which provide a supporting role for the guard cells.
Stomata begin as stomatal meristemoids. The process varies between dicots and monocots. Spacing
is thought to be essentially random in dicots though mutants do show it is under some form of genetic
control, but it is more controlled in monocots, where stomata arise from specific asymmetric
divisions of protodermal cells. The smaller of the two cells produced becomes the guard mother cells.
Adjacent epidermal cells will also divide asymmetrically to form the subsidiary cells.
Because stomata play such an important role in the plants survival, collecting information on their
differentiation is difficult by the traditional means of genetic manipulation, as stomatal mutants tend to
be unable to survive. Thus the control of the process is not well understood. Some genes have been
identified. TMM is thought to control the timing of stomatal initiation specification andFLP is thought to
be involved in preventing further division of the guard cells once they are formed.
Environmental conditions affect the development of stomata, in particular their density on the leaf
surface. It is thought that plant hormones, such as ethylene and cytokines, control the stomatas
developmental response to the environmental conditions. Accumulation of these hormones appears
to cause increased stomatal density such as when the plants are kept in closed environments.
Stomatal cells only occur on the leaf epidermis, and it is thought that inhibitory signals must occur on
other parts of the plants epidermis to prevent stomatal formation there. These signals could be
hormonal, or perhaps gene products transmitted from underlying tissues via the plasmodesmata


Why onion epidermal cells do not contain
Chloroplast?

They don't need it.
An onion plant has two parts- 1. Onion bulbs (the part which is pink in color) and 2.onion leaves.
Onion bulbs grow 'under' the soil while onion leaves are slender and straight and grow above the
ground towards the sunlight just like other plants do. The onion 'leaves' contain chloroplasts not the
'bulbs'.
You can see in the following picture how onion leaves look like and remember that the onion bulbs
are growing just underneath the soil.
Do the epidermal cells of a leaf contain chloroplast?

No, regular epidermal cells do not have chloroplasts. The guard cells that surround the stoma do have
chloroplasts, but they are not the regular epidermal cells.

Why doesn't the upper epidermis of the leaf have
chloroplasts?
The epidermis serves several functions: protection against water loss, regulation of gas exchange
(although, this is mostly served at the lower epidermis), secretion of metabolic compounds, and (in
some species) absorption of water.

The epidermis is usually transparent (as you said, epidermal cells lack chloroplasts) and coated on
the outer side with a waxy cuticle that prevents water loss.

Epidermis
The epidermis is the outermost cellular layer which covers the whole plant
structure, i.e. it covers roots, stem, leaves, flowers and fruit. It is composed of a
single layer of living cells, although there are exceptions. Epidermis is usually
closely packed, without intercellular spaces or chloroplasts. The outer walls, which
are exposed to the atmosphere and usually thickened, and may be covered by
a waxy, waterproof cuticle which are made up ofcutin. Apart from the normal
epidermal cells there are also stomata in the epidermis of leaves and stem.
A stoma is an opening (pore) which is bounded by two beanshaped cells
called guard cells . The guard cells differ from normal epidermal cells in that they
have chloroplasts and the cell walls are thickening unevenly; the outer wall is thin
and the inner wall (nearest the opening) is thick. The thin-walled epidermal cells of
roots give rise toroot hairs. Hair- like outgrowths may also be found in the
epidermis of leaves and stems.
Functions:
o the epidermal cells protect the underlying cells,
o the waxy cuticle prevents the loss of moisture from the leaves and stems,
o the transparent epidermal cells allow sunlight (for photosynthesis) to pass
through to the chloroplasts in the mesophyll tissue,
o the stomata of leaves and stems allow gaseous exchange to take place
which is necessary for photosynthesis and respiration,
o water vapour may be given off through the stomata during transpiration,
o the root-hairs absorb water and dissolved ions from the soil
Root epidermis not covered with cuticle
Roots take in water and the stem tries to
conserve water. That simple.
o The cuticle is water proof - the root epidermis
eneeds to be semi-permiable for
o osmosis to occur

Epidermis
The root hairs of the young epidermal cells vastly increase the surface area through
which movement of materials can occur. The thread-like hairs are simply enlargements
of the protoplast that extend outward into the soil. They have little wall material and are
extremely fragile and easily broken. The root epidermis of some plants is covered by a
thin, waxy cuticle, which apparently isn't thick enough to impede movement of
substances through the epidermis.
Cortex
The cortex, composed primarily of parenchyma cells, is the largest part of the primary
root, but in most dicots (eudicots) and in gymnosperms that undergo extensive
secondary growth, it is soon crushed, and its storage function assumed by other tissues.
Three layers of cortex are recognized: the hypodermis (also called exodermis),
the endodermis and, between them, the storage parenchyma. The outer and inner
layers of the cortex, the hypodermis and endodermis, are cylinders of tightly packed
cells with heavily suberized walls and no intercellular spaces. (Suberin is the fatty
substance that gives cork its distinctive attributes.) In contrast, the storage parenchyma
cells are thin-walled and loosely packed with many intercellular spaces among them.
Hypodermis (exodermis). Just under the epidermis forming the outermost layer of the
cortex is a layer one or two cells in width called the hypodermis. Since its cell walls are
heavily suberized and impermeable to water its apparent function is to keep the water
and nutrients (which are absorbed in the root zone further down the root) from leaking
out through the cortex. The hypodermis is especially well developed in plants of arid
regions and in those with shallow root systems. It also deters the entrance of soil
microorganisms.
Endodermis. The innermost layer of the cortex is the endodermis, which is readily
identifiable by the presence of Casparian strips, bands of suberin present on
transverse and radial walls of its cellsthe walls perpendicular to the surface of the root.
The endodermis regulates the passage of water and dissolved substances by forcing
them to move through living plasma membranes and plasmodesmata and not simply
diffuse through the porous cell walls. The absorption and translocation of materials is
thus selective; not everything in the surrounding soil gets through and into the plant
body. An endodermis almost always is present in roots and generally never in stems.
Storage parenchyma. The bulk of the cortex consists of thin-walled, living parenchyma
cells, which store starch and other substances. The cells expand or shrink as materials
move in and out of their protoplasts. The large volume of air present in the intercellular
spaces of this tissue provides important aeration for roots.
Stele (vascular cylinder)
The stele includes all of the tissues inside of the cortex: the pericycle, the vascular
tissuesxylem and phloemand, in some plants, a pith. Most dicot (eudicot) roots have
a solid core of xylem in their center whereas most monocots have a pith composed of
parenchyma.
Pericycle. The pericycle is a cylinder of parenchyma, one or at most a few cells in width,
which lies in the stele immediately inside the endodermis. The cells retain their ability to
divide throughout their lives, and localized divisions in the pericycle give rise to lateral
(branch) roots. When secondary growth occurs in roots, the vascular cambium and
usually the first cork cambium originate in the pericycle. Other cell divisions in the
pericycle produce additional pericycle cells.
Vascular tissues. Most dicot (eudicot) roots differ from eudicot stems in having a lobed
column of primary xylem as their core with phloem tissue occurring as strings of cells
between the lobes. This arrangement is called a protostele. The primary xylem of
monocots, on the other hand, forms a cylinder around a central mass of pith
parenchyma, a siphonostele. The way in which the vascular tissues develop is useful in
tracing ancestral relationships in the plant kingdom.

Taiz and Zeiger on line
Topic 1.4
Plant Tissue Systems: Dermal, Ground, and Vascular
Dermal Tissue
The epidermis is the dermal tissue of young plants
undergoing primary growth (see textbook Figure 1.3). It is
generally composed of specialized, flattened polygonal cells
that occur on all plant surfaces. Shoot surfaces are usually
coated with a waxy cuticle to prevent water loss and are
often covered with hairs, or trichomes, which are epidermal
cell extensions.
Pairs of specialized epidermal cells, the guard cells, are
found surrounding microscopic pores in all leaves (see
textbook Figure 1.3A). The guard cells and pores are
called stomata (singular stoma), and they permit gas
exchange (water loss, CO
2
uptake, and O
2
release or uptake)
between the atmosphere and the interior of the leaf.
The root epidermis is adapted for absorption of water and
minerals, and its outer wall surface typically does not have a
waxy cuticle. Extensions from the root epidermal cells,
the root hairs , increase the surface area over which
absorption can take place (see textbook Figure 1.3C).
Ground tissue. Making up the bulk of the plant are cells
termed the ground tissue. There are three types of ground
tissue: parenchyma, collenchyma, and sclerenchyma.
Parenchyma, the most abundant ground tissue,
consists of thin-walled, metabolically active cells that
carry out a variety of functions in the plant, including
photosynthesis and storage (see textbook Figure 1.3B)
Collenchyma tissue is composed of narrow, elongated
cells with thick primary walls (see textbook Figure
1.3C). Collenchyma cells provide structural support to
the growing plant body, particularly shoots, and their
thickened walls are nonlignified, so they can stretch as
the organ elongates. Collenchyma cells are typically
arranged in bundles or layers near the periphery of
stems or leaf petioles.
Sclerenchyma consists of two types of cells, sclereids
and fibers (see textbook Figure 1.3D) Both have thick
secondary walls and are frequently dead at maturity.
Sclereids occur in a variety of shapes, ranging from
roughly spherical to branched, and are widely
distributed throughout the plant. In contrast, fibers are
narrow, elongated cells that are commonly associated
with vascular tissues. The main function of
sclerenchyma is to provide mechanical support,
particularly to parts of the plant that are no longer
elongating.
In the stem, the pith and the cortex make up the ground
tissue (see textbook Figure 1.1B). The pith is located within
the cylinder of vascular tissue, where it often exhibits a
spongy texture because of the presence of large intercellular
air spaces. If the growth of the pith fails to keep up with that
of the surrounding tissues, the pith may degenerate,
producing a hollow stem. In general, roots lack piths,
although there are exceptions to this rule. In contrast,
the cortex , which is located between the epidermis and the
vascular cylinder, is present in both stems and roots (see
textbook Figure 1.1B and C).
At the boundary between the ground tissue and the vascular
tissue in roots, and occasionally in stems, is a specialized
layer of co rtex known as the endodermis (see textbook
Figure 1.1C). This single layer of cells originates from cortical
tissue at the innermost layer of the root cortex and forms a
cylinder that surrounds the central vascular tissue, or stele.
Early in root development, a narrow band composed of the
waxy substance suberin is formed in the cell walls
circumscribing each endodermal cell (see textbook Figure
1.1). These suberin deposits, called Casparian strips, form
a barrier in the endodermal walls to the intercellular
movement of water, ions, and other water-soluble solutes to
the vascular cells.
Leaves have two interior layers of ground tissue that are
collectively known as the mesophyll (see textbook Figure
1.1A). The palisade parenchyma consists of closely
spaced, columnar cells located beneath the upper epidermis.
There is usually one layer of palisade parenchyma in the leaf.
Palisade parenchyma cells are rich in chloroplasts and are a
primary site of photosynthesis in the leaf. Below the palisade
parenchyma are i rregularly shaped, widely spaced spongy
mesophyll cells. The spongy mesophyll cells are also
photosynthetic, and the large spaces between these cells
allow diffusion of carbon dioxide. The spongy mesophyll also
contributes to leaf flexibility in the wind, and this flexibility
facilitates the movement of gases within the leaf.
Vascular tissues: xylem and phloem. The vascular tissue
is composed of two major conducting systems: the xylem
and the phloem. The xylem transports water and mineral
ions from the root to the rest of the plant.
The phloem distributes the products of photosynthesis and a
variety of other solutes throughout the plant (see textbook
Figure 1.1B and C).
The tracheids and vessel elements are the conducting
cells of the xylem (see textbook Figure 1.3E). Both of these
cell types have elaborate secondary-wall thickenings and lose
their cytoplasm at maturity; that is, they are dead when
functional. Tracheids overlap each other, whereas vessel
elements have open end walls and are arranged end to end
to form a larger unit called a vessel. Other cell types present
in the xylem include parenchyma cells, which are important
for the storage of energy-rich molecules and phenolic
compounds, and sclerenchyma fibers.
The sieve elements and sieve cells are responsible for
sugar translocation in the phloem (see textbook Figure 1.3E).
The former are found in angiosperms; the latter perform the
same function in gymnosperms. Like vessel elements, sieve
elements are often stacked in vertical rows, forming larger
units called sieve tubes, whereas sieve cells form
overlapping arrays. Both types of conducting cells are living
when functional, but they lack nuclei and central vacuoles
and have relatively few cytoplasmic organelles.
Substances are translocated from sieve cell to sieve cell
laterally through circular or oval zones containing enlarged
pores, called sieve areas. In contrast, sieve tubes
translocate substances through large pores in the end walls
of the sieve elements, called sieve plates. Sugar movement
through sieve tubes is more efficient and rapid than through
sieve cells and represents a more evolutionarily advanced
mechanism.
Sieve elements are associated with, and depend on, densely
cytoplasmic parenchyma cells calledcompanion cells. The
analogous cells adjacent to the sieve cells of gymnosperms
are called albuminous cells. Companion cells provide
proteins and metabolites necessary for the functions of the
sieve tube elements. In addition, the phloem frequently
contains storage parenchyma and fibers that provide
mechanical support.
Chapter 46 - Plant Structure
Monocots Vs Dicots
Monocots Dicots
Number of Cotyledons one two
Vascular Tissue - Roots arranged in a ring phloem between arms of
xylem
Vascular Tissue - Stems bundles scattered bundles form a ring
Veins in Leaves parallel net-like pattern
Number of Flower Parts 3 or multiples of 3 4 or 5 or multiples of 4 or 5

Roots and Shoots
Roots anchor, absorb H
2
O and minerals, and store starch.
The shoot is the above-ground portion of the plant. The stem transports water and
minerals to the leaves and sugar to the roots.

Tissue types
Plants have four tissue types.
Vascular tissue transports, dermal tissue protects, meristematic tissue grows (cells
divide), and ground tissue forms the rest of the plant.

Vascular tissue
Xylem
transports water and minerals from roots to leaves
composed of hollow, nonliving cells
tracheids- elongated with tapered ends, pits or depressions
vessel elements- larger, forms a continuous pipeline
Phloem
Transports organic nutrients, usually from leaves to roots
Phloem cells are living.
sieve-tube cells- no nucleus; connected to each other by plasmodesmata
Sieve plates are found at the ends of the cells.
Companion cells contain a nucleus and are located in close proximity to sieve-tube
cells. They are connected to sieve tube cells by plasmodesmata.

Meristematic tissue
Areas within the plant that are capable of growth (cell division) are
called meristems.
Primary Growth
Primary growth occurs only at the shoot and root tips in areas called apical
meristems. Primary growth is responsible for elongating the plant. In areas that
contain only primary growth, stem thickness increases by cell enlargement, not by
the production of new cells.
Secondary Growth
Lateral meristems produce new cells that make the stems and roots thicker. This
type of growth is called secondary growth. Secondary growth occurs only during
the second and subsequent years and only in woody species.
There are two kinds of lateral meristems, the vascular cambium and the cork
cambium. These lateral meristems form as rings within the plant body as the stem
increases in thickness. The diagrams below illustrate how the vascular cambium
divides to produce new xylem cells toward the inside of the vascular cambium and
new phloem cells toward the outside.

The vascular cambium cells divide longitudinally.

One of the new cells remains vascular cambium and the
other becomes xylem. The cells can be seen enlarging in
this diagram.

Occasionally,
the inner cells remain vascular cambium and the outer ones become
phloem.

Dermal tissue
outer covering of plant
It consists of closely packed cells that function to protect.
The epidermis covers the plant but is replaced by cork (periderm) in the stems and
roots of woody plants.
The epidermis has a waxy covering called a cuticle that protects the plant from
desiccation.
The periderm is the outer part of the bark.

Ground tissue
Ground tissue fills interior of plant. It contains parenchyma, collenchyma, and
sclerenchyma cells.
Parenchyma
thin-walled
least specialized of the three cell types
found in all organs
usually functions in photosynthesis or storage
photosynthetic parenchyma have chloroplasts
parenchyma that function for storage have colorless plastids
Parenchyma can divide to produce more specialized types of cells.
Collenchyma
Collenchyma cells have thicker primary cell walls, especially at the corners.
A primary cell wall is one that is produced while the cell is growing.
Collenchyma often forms bundles just beneath the epidermis for flexible support of
immature parts of the plant body.
Sclerenchyma
Sclerenchyma cells have thick secondary cell walls, usually toughened with lignin.
A secondary cell wall is one that is produced after the cell is mature. It is produced
inside the primary cell wall.
Most sclerenchyma cells are nonliving. They function to support mature regions
and produce hard parts (example: nut shells).
omit: contain fibers (long and slender) and sclereids (shorter, varied shape)
example: sclereids make nut shells hard
Primary growth of stem
Primary growth occurs only in apical meristems which are located at the tips of the
stems and roots.
Meristems in stems are protected by newly formed leaves within a bud.

Axillary buds
usually dormant
in the axes of mature leaves
develop into branches

Herbaceous stems (nonwoody
Herbaceous stems are produced by primary growth.
The outermost tissue is epidermis and is covered by waxy cuticle to prevent water
loss.
The vascular tissue is found in bundles that are arranged in a ring (dicots) or
scattered (monocots).


In dicots, the xylem is toward inside; the phloem is toward the outside.
cortex- In dicot stems, the cortex is located in the area between the vascular
bundles and the epidermis. In monocot stems, it occupies the area surrounding the
vascular bundles.
The center of the stem is pith and may function as storage.

Secondary Growth of Stems
Secondary growth occurs in plants that live > 1 year.
primary growth occurs for a short distance behind the apical meristem,
then secondary growth occurs.
It begins with the formation of a vascular cambium and a cork cambium.


ebVascular cambium
Initially, vascular cambium is found between the xylem and phloem in the vascular
bundles of dicots.
After one years growth, it joins to form a continuous ring.
Cell division toward the inside and outside form xylem and phloem.
Seasonal climates produces growth rings because cells grow faster and are larger in
the spring than later in the growing season.



Cork cambium
Cortex cells beneath the epidermis produce the cork cambium.
The cork cambium produces cork.
Cork is waterproof because the cell walls are impregnated with of suberin.
Pockets of cells lack suberin. These are called lenticels and function to allow gas
exchange.
Cork replaces the epidermis on woody stems and roots.

Bark
The bark of trees consists of cork, cork cambium, cortex, and phloem.

Summary of Stem Growth
Primary Growth Lateral Meristems Secondary Growth
Dermal Tissue epidermis cork
Ground Tissue cortex
pith

Meristem Tissue vascular cambium cork cambium
vascular cambium

Vascular Tissue primary phloem
primary xylem
secondary xylem and
phloem

Stem External Structure
Stems support, conduct, store water and photosynthate (products of
photosynthesis).
nodes- where leaves attach
internodes - between nodes
bud - contains apical meristem and newly-forming leaves
leaves - photosynthetic organs

Types of stems
stolons (runners)- horizontal, aboveground- strawberries
rhizomes- horizontal, underground; responsible for rapid spread of many weeds
tubers- enlarged tips of rhizomes; food storage- potato
corms- underground, short, thick, vertical; food storage- gladiolus
bulbs- underground with thick, fleshy leaves- onion
tendrils- assist plant in climbing

Leaves
Leaves usually function in photosynthesis, so they are flattened to increase the
surface exposed to light.
blade, petiole
simple, compound
pinnate, palmate
opposite,alternate, whorled
vary according to environmental conditions
broad in shade
reduced in dry areas (ex: spines in cacti)
succulent leaves hold water
can be adapted for food storage (onions)
climbing leaves can be modified as tendrils
Monocot leaves have parallel veins; dicot leaves have a net-like pattern.
The top layer is the epidermis, a type of dermal tissue. It often has protective hairs
and/or glands that produce irritants.
always a waxy cuticle
mesophyll: parenchyma cells w/ chloroplasts
Stomata
Stomata (sing. stoma) are openings in the epidermis of leaves and stems that allow
gas exchange. Guard cells surround the opening and function to open or close
it. Guard cells that contain chloroplasts, other epidermal cells do not contain
chloroplasts
When K
+
is pumped into the guard cell by active transport (requires ATP), water
follows by osmosis. This causes the cells to bend and open. When K
+
(and water)
leaves the guard cells, they close.

C
3
plants
palisade layer and spongy layer
The loss of water from the leaves by evaporation is called transpiration. It
accounts for more than 90% of water taken up by the roots.


Dicot roots
zone of cell division- root apical meristem; just behind the root cap
zone of elongation- area where cells elongate; become more specialized
zone of maturation (differentiation)
cells mature and become fully differentiated (specialized)
The epidermal cells form root hairs in this zone. Root hairs increase the absorptive
surface area.

Specialized tissues of roots
Epidermis
outer layer consisting of rectangular-shaped cells
Root hairs are extensions of epidermal cells that project 5-8 mm into the soil. They
increase the surface area of the root for absorption.
Cortex
interior to epidermis
large thin-walled parenchyma; loosely packed
water can move through cortex without entering cells
starch granules in cortex function for storage
Vascular tissue
xylem- star-shaped
phloem- between rays of xylem
Endodermis
The endodermis is a single layer of cells that forms a boundary between the cortex
and the inner vascular cylinder.
The endodermis is lined on 4 sides by the Casparian strip. The casparian strip is a
coating that prevents water from seeping between the cells and thus forces water to
enter the endodermal cells before passing through to the vascular cylinder.



The Casparian strip surrounds cells of the
endodermis and prevents water and
minerals from seeping between the cells.
In order to get to the vascular cylinder,
water and minerals must pass through
the cell membrane.
Without the casparian strip, water and
minerals would be able to enter the
vascular cylinder by going between cells.
Pericycle
The pericycle is the layer just inside the endodermis.
It retains the capacity to divide and form branch roots.

Monocot roots
Unlike monocot stems, the vascular tissue in monocot roots is arranged in a ring.
Monocot roots are like dicot roots in that they contain pericycle, endododermis,
cortex (outside of vascular tissue), and an epidermis.
The central portion of the root is called pith. It is composed of parenchyma and
functions in storage.
Monocot roots typically have no secondary growth.

Secondary growth in roots
Secondary growth in roots is similar to stems; annual growth rings are formed.
Vascular cambium forms between xylem and phloem.
The pericycle produces the cork cambium.


Primary Growth in a dicot root.




As the root increases in diameter, the vascular cambium
becomes circular.


Summary of Root Growth
Primary Growth Lateral Meristems Secondary Growth
Dermal Tissue epidermis cork
Ground Tissue cortex
pith


Meristem Tissue
pericycle
vascular cambium
cork cambium
vascular cambium

Vascular Tissue primary xylem and phloem secondary xylem and
phloem

Root Systems
Dicots
the primary (first) root grows straight down; called a taproot
often fleshy and stores food; ex: carrots, beets, turnips, radishes

Monocots
fibrous root system, no main root
Adventitious roots- new roots that arise from an aboveground structure; example-
prop roots on corn


Stems Roots



Primary
Growth
(Monocots)




Primary
Growth
(Dicots)




Secondary
Growth
(Dicots)


A plant cuticle is a protective film covering the epidermis of leaves, young shoots and other
aerial plant organs without periderm. It consists of lipid and hydrocarbon polymers impregnated with
wax, and is synthesized exclusively by the epidermal cells.
[1]
A cuticle is present in
the sporophytegeneration of hornworts, in both sporophyte and gametophyte generations
of mosses
[2]
and in the sporophytes of all vascular plants. Inangiosperms the cuticle tends to be
thicker on the top of the leaf (adaxial surface), but is not always thicker in xerophytic plants living in
dry climates than in mesophytic plants from wetter climates, despite a persistent myth to that effect.
The cuticle is composed of an insoluble cuticular membrane impregnated by and covered with
soluble waxes. Cutin, a polyester polymercomposed of inter-esterified omega hydroxy acids which are
cross-linked by ester and epoxide bonds, is the best-known structural component of the cuticular
membrane.
[3][4]
The cuticle can also contain a non-saponifiable hydrocarbon polymer known
as Cutan.
[5]
The cuticular membrane is impregnated with cuticular waxes
[6]
and covered
with epicuticular waxes, which are mixtures of hydrophobic aliphatic compounds, hydrocarbons with
chain lengths typically in the range C16 to C36.
[7]

The plant cuticle is one of a series of innovations, together with stomata, xylem and phloem and
intercellular spaces in stem and later leafmesophyll tissue, that plants evolved more than 450 million
years ago during the transition between life in water and life on land.
[8]
Together, these features
enabled upright plant shoots exploring aerial environments to conserve water by internalising the gas
exchange surfaces, enclosing them in a waterproof membrane and providing a variable-aperture
control mechanism, the stomatal guard cells, which regulate the rates of transpiration and
CO
2
exchange.
In addition to its function as a permeability barrier for water and other molecules, the micro and nano-
structure of the cuticle confer specialised surface properties that prevent contamination of plant
tissues with external water, dirt and microorganisms. Aerial organs of many plants, such as the leaves
of the sacred lotus (Nelumbo nucifera) have ultra-hydrophobic and self-cleaning properties that have
been described by Barthlott and Neinhuis (1997).
[9]
The lotus effect has uses in biomimetic technical
materials.
"The waxy sheet of cuticle also functions in defense, forming a physical barrier that resists penetration
by virus particles, bacterial cells, and the spores or growing filaments of fungi".


DO NOT PRINT THIS PAGE. Click HERE for your
print-friendly copy.

The Root
This organ is the first to emerge from a seed.
Its functions are
anchor the plant to its substrate
absorb water and inorganic substances
from the substrate
conduct the above upwards to the rest of
the plant
production (in meristems) of certain
hormones that are transported other
parts of the plant
production of secondary metabolites (e.g.,
nicotine in tobacco, which is transported
to the leaves for deposition as an
herbivore deterrent)
storage of nutrients as carbohydrates
and/or lipids
Origin of the Root
The primary root is the first root that forms in
and emerges from the seed.
In most other plants, the primary root develops
into a taproot, a large, central root from
which lateral roots emerge.

Root depth depends on environment
Porous soil - deep roots; rocky soil - shallower roots
Temperature and soil water content also affect root depth.
And of course, different species have different root characteristics
spruce, beech, poplar, tend to have relatively shallow roots
oaks, most pines, maples, sycamores, etc, tend to have taproots almost as tall as the
aboveground portion of the plant.
What about southern Florida root depth? The dilemma of hurricanes!
record root depth is held by the mesquite: just over 53 METERS. (Why mesquite?)
The main "feeder roots" spread in the topmost soil layers, usually no deeper than about a meter.
Roots tremendously increase plant surface area where the plant needs it: for water absorption.
Root surface area is much greater than shoot surface area.
Root and shoot are balanced: If the roots are damaged, the shoot may die back.
If the shoot is damaged, there's less photosynthate available for new root growth. In the most
derived plants (monocots), the taproot is replaced by a system of fibrous roots that all emerge in a
bunch at the base of the stem.



Root Growth and Development
Let's have a look at where it all begins: The Root Tip.
Again, note the relative locations of the apical and three primary meristems.


Can you see the quiescent center? Labeled with a radioactive nucleoide, the tip shows up on an
autoradiograph with actively dividing nuclei showing dark (taking up the marker), and relatively
inactive cells with invisible nuclei (not taking up marker).
The meristematic quiescent center
gives rise to the root cap cells, replacing them as the old ones are sloughed off
organizes the growth pattern of the three primary meristems which also arise from it
Root Cap
The root cap is a protective cap of live parenchyma cells
it is produced by the apical meristem behind it
it protects the root meristem as it grows through the soil
outermost cells are sloughed off as the root tip "pushes" through the soil
root cap cells produce a slimy lubricant (mucilage or mucigel)
o it's the plant equivalent/analog of mucus
o hydrophilic polysaccharide (a type of pectin)
o also contains sugars, organic acids, vitamins, enzymes, and amino acids.
o produced in the Golgi and packaged in vesicles that attach to the cell wall and empty
their contents.
o the goo passes through the cell wall and oozes out onto the root cap surface.
o mucigel allows greater ease of transporting ions from interstitial soil water to the
root surface, where it can be absorbed.
o mucigel is friendly to nitrogien-bacteria, and may help them colonize roots of
nitrogen-fixing plant species (primarily in the Pea Family, Fabaceae)
o may provide protection against desiccation.
o in some plants, mucigel contains inhibitors that prevent the growth of roots from
competing plants. (allelopathy)
o aids in water and nutrient absorption by increasing soil/root contact.
o mucigel can chelate soil ions, making them available to the root.
o mucigel components can aid in the establishment of mycorrhizae and symbiotic
bacteria.
The root cap cells last 4-9 days, depending on plant species and growth rate
before they become root cap cells, root cap meristem cells differentiate into the cells of
the columella a central column of cells that contain starchy amyloplasts.
In response to gravity, the amyloplasts fall to the bottom of the cells, attracting hormones
that promote growth in the direction of the amyloplasts.
Collumella cells are also light-sensitive, and respond to pressure from soil particles, further
signalling to the plant which way is down.
By the time columella cells are pushed to the periphery of the root cap by other developing
cells behind them, they have differentiated into peripheral root cap cells.
Peripheral root cap cells secrete mucigel that is manufactured in their dictyosomes
(modified Golgi apparatus).

Root Meristems
The apical meristem (from the Greek merismos, which means "division") is found at the very tip of
the root, just behind the root cap.
As in all meristems, the apical meristem contains some cells that will always remain meristematic:
one daughter cell remains in the meristem (the initial) and continues to divide, whereas its sister cell
(the derivative) stays behind as the meristem grows out. The derivative differentiates into some type
of cell, depending on its gene expression. This is known as primary growth.
The very end of the root tip contains the initials and the immediate derivatives, and is known
as the promeristem. Note the relative locations of the apical and three primary meristems.

(What's different about this quiescent center? Why?)
Region of Cell Division - the apical meristem
Region of Elongation - growing cells elongating (primary meristems)
Region of Maturation - cells mature, no longer elongating (mature tissues)

Root Primary Structure
Recall that any plant organ has three main layers:
epidermis (and derivatives)
vascular tissue
ground tissue

Root Anatomy: Cross Sectional View

From outermost layer to innermost:
epidermis
cortex
endodermis - selectively permeable layer; unique to roots
pericycle - a secondary/lateral meristem that gives rise only to side branch roots; unique to
roots
vascular cylinder (a.k.a. stele)

Root Epidermis
Root epidermis is the surface that meets the environment, and it is the first selectively permeable
membrane the plant uses to filter uptake.
Surface area is increased by trichomes that form root hairs:

These are found primarily in the Region of Maturation, and die off once the cells age.
Although the cell walls contain suberin, water and minerals can pass easily between the cells
of the epidermis, so further filtration is needed down the line.

Exam I material ends here.
Exam II material begins here.

Mycorrhizae
This is a symbiotic relationship between a fungus and a plant root. (What does each partner get out
of the relationship?)
Vesicular Arbuscular Mycorrhizae (V.A.M.) - association between a zygomycete fungus
("Black Bread Mold") and a plant
Ectomycorrhizae - association between ascomycete (Sac Fungus) or basidiomycete (Club
Fungus) and a conifer or flowering plant (usually large trees).

Some of the most valuable edible organisms in the world are TRUFFLES, various species of
mycorrhizal (ascomycete and basidiomycete) fungi that partner with plants.
In mycorrhizal plants, root hair surface area is negligible compared to that provided by the
interface of mycorrhiza, plant and fungus. Most absorption is done via the mycorrhizal
hyphae.
Recent research suggests that mycorhizzal associations may be a symbiotic partnership
between not two, but three species, including special bacteria that live inside the gungus and
are essential to the establishment of the symbiosis.

The Cortex
Just internal to the epidermis lies the cortex, composed primarily of parenchyma.

Cortex plastids are primarily for storage (fats, carbs). Only in some species with
photosynthetic roots (which types of plants would you expect these to be?) are there
chloroplasts in these cells.
In woody plants, the cortex is shed off once woody growth begins. In herbaceous plants, the
cortex is maintained throughout the life of the plant.
Most of the cortex is airy, with a lot of space (filled with fluid and or air) between the cells.
Fluids travel via:
symplast - the connection formed by plasmodesmata
apoplast - the continuous surface formed by adjoining cell walls
(recall: the tonoplast is the continuous fluid pathway formed by the plasma membrane of the
vacuoles)
The innermost layer of the cortex is the endodermis, the main "filtration" surface of the root.

The Casparian strips banding each endodermal cell are made of suberin (sometimes lignin,
as well), and prevent interstitial entry of water into the stele (central core of vascular tissue).
Thus, water cannot travel via the apoplast, and must pass through the selectively permeable
plasma membrane of the endodermal cells before it reaches the vascular system.

Pericycle
This is a layer of pluripotent parenchyma cells located just inside the endodermis. Pericycle gives rise
to side branch roots.


The Stele
Root morphology is fairly well conserved across plant taxa. Therefore, differences in the morphology
of the stele can be an important tool for classifying plants and determining evolutionary relatedness.
The stele consists (from outermost to innermost layers):
phloem from three to many bundles, alternating with xylem
xylem - central core
The number of xylem "arms" in the stele determines its classification as a...
o diarch - two arms (uncommon)
o triarch - three arms
o tetrarch - four arms
o pentarch - five arms
o hexarch - six arms
o polyarch - more than six arms ...stele.

In young roots, the phloem and xylem are nested in a central core of parenchymal pith
In monocot roots only the xylem core surrounds a central cylinder of parenchymal pith. This
is the simplest way to tell the difference between a typical root and the highly derived
monocot root.

The Variety of Roots
Roots of various plant species have evolved various specializations.
food storage roots - used by the plant to store starch for metabolic activities later in the
season. Typical examples: carrot, beet, sweet potato.
water storage roots - found in arid regions, these are roots that collect large amounts of
water during rainy season for the plant to use during dry season. These are most often
found in xeriphytes (sometimes spelled xerophyte). Local examples include the East Indian
Rosewood and the Starburst.
propagative roots - have meristematic regions from where new, genetically identical
plantlets can grow. These regions are not the same as nodes: they do not contain a true
apical meristem.
Local examples include the East Indian Rosewood and the Starburst.
pneumatophores - gas exchange surfaces on root tips protruding from water-logged soil.
Certain species of mangrove have these. But, contrary to popular myth, cypress
"knees" apparently have no gas exchange function. (Cypress trees with knees removed do
not suffer from any apparent lack of oxygen.
prop roots - These grow from the lower part of a stem or trunk down to the ground, and
providing extra support for the plant. These tend to be more common in plants with a tall,
soft stem structure, as well as in plants that live in softer soils.
Common examples include corn (Zea mays), Screw "Pine" (Pandanus tectorius), various
species of palms, and red mangroves (Rhizophorus mangle.
aerial roots - typical of epiphytes such as orchids (in which these roots are called velamen,
with a spongy outer surface very good at absorbing and holding water) and bromeliads.
buttress roots - wall-like extensions off the base of the trunk which provide support against
physical assault from high winds.
Our local Ficus spp. and the Royal Poinciana (Delonix regia tend to develop these in certain
environments.
contractile roots - these specialized roots, usually found at the base of an underground
organ (e.g., a bulb) actually contract to perform such functions as getting a bulb to its proper
soil dept for growth
haustoria - parasitic plant roots that invade the tissues of a host plant and transfer nutrients
from host to parasite.
Examples of plants that have haustoria are dodder and mistletoe
adventitious roots - are roots that grow anywhere they are not "expected." Examples are
the adventitious roots that grow so prodigiously from some of our native and introduced
species of Ficus trees. Several of the root types listed above (e.g., prop roots, aerial roots)
can also be considered adventitious.

Life-sustaining Root Symbiosis: Nitrogen Fixation
The Nitrogen Cycle is the pathway by which nitrogen moves through living and non-living
components of the ecosystem.
Nitrogen is one of the four main elements most common in biological macromolecules, and
yet no eukaryotes are capable of fixing atmospheric nitrogen , N
2
, into its usable forms, such
as ammonium (NH
4
+
) with other species changing it into nitrite (NO
2
-
) and nitrate (NO
3
-
).
Certain nitrogen-fixing bacteria, however, are capable of converting gaseous nitrogen into its
biologically useful forms, and some of these have formed symbiotic relationships with plants,
notably in the Fabaceae (Pea Family), commonly called legumes.
The roots of legumes are covered with swellings called nodules within which reside
symbiotic bacteria that fix nitrogen. Various strains of a bacterial species
named Rhizobium form this association.
Nitrogen fixation into ammonium requires an anaerobic environment such as that found in
the root nodules. The root nodule surfaces are highly lignified, helping to prevent gas
exchange. Also, root nodules often contain leghemoglobin, a hemoglobin-like molecule with
high affinity for free oxygen. This protein provides a sort of "buffer" for oxygen, allowing the
bacteria enough oxygen to produce ATP for the very energy-expensive reactions of nitrogen
fixation without allowing too much oxygen to build up in the nodule tissues and interfere
with nitrogen fixation itself.
The figure below shows the sequence of events leading to nodule formation.

How does this symbiosis develop? It's amazing...
The plant root emits flavonoids into the soil.
Certain species of Rhizobium take up these flavonoids (the strain of Rhizobium colonizing
each plant species is different, and determined by the exact structure of the flavonoid
messenger.)
The flavonoid activates a transcription factor protein, the activity of which results in the
activation of a bacterial operon known as nod(for "nodule").
The genes in the nod group produce enzymes that catalyze Nod proteins, specific to the
bacterial strain.
The Rhizobium secrete the Nod molecules into the soil, and these signal to the plant root to
elongate root hairs and form the infection thread that the bacteria will use to enter the
root.
There is some evidence to suggest that early mycorrhizal fungus/plant communication
pathways (which also employ flavonoids) led to the evolution of the bacteria/plant
communications resulting in nitrogen fixation symbiosis.
Crop Rotation
Most agricultural crops severely deplete soil nitrogen. Hence, good farming technique usually
includes crop rotation, in which the farmer will grow a non-legume crop in a field for one or more
years, and then plants a legume crop for a year to help restore soil fixed nitrogen.