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 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 of mucus
 o hydrophilic polysaccharide (a type of pectin)
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 some protection against desiccation.
 root cap cells last 4-9 days, depending on plant species and growth rate
 root cap facilitates geotropism via the columella, a central column of cells that contain
starchy amyloplasts. These fall to the bottom of the cells, attracting hormones that
promote growth in the direction of the amyloplasts. (more on this later)

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.

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 Nitrogen is one of the four main
elements most common in biological macromolecules, and yet no eukaryotes are capable
of fixing atmospheric nitrogen , N2, into its usable forms, such as ammonium (NH4+) with
other species changing it into nitrite (NO2-) and nitrate (NO3-).

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...

o The plant root emits flavonoids into the soil.

o 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.)
o The flavonoid activates a transcription factor protein, the activity of which results
in the activation of a bacterial operon known as nod (for "nodule").
o The genes in the nod group produce enzymes that catalyze Nod proteins, specific
to the bacterial strain.
o 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.