Epidermis (botany)
Encyclopedia
The epidermis 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.
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 or long filaments. Thick 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. 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.
Trichome
s 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
.
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 number of stomata varies from about 1,000 to over 100,000 per square centimeter of leaf surface.
The stoma is bounded by two guard cells. The guard cells differ from the epidermal cells in the following aspects:
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.
s and their subsidiary cells that surround the stomata and trichome
s, 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 gene
s: TTG and GL1. The process may be controlled by the plant hormone
s gibberellin
s, 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 petal
s of flower
s. 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 lot 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 mutant
s 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 and FLP 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 stomata’s 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
.
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
, flowers, roots and stems
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
. It forms a boundary between the plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
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.
Description
The epidermis is the outermost cell layer of the primary plant body, it is the dermal tissue system of leaves (diagrammed below), stems, roots, flowers, fruits, and seeds; it is usually transparent (epidermal cells lack chloroplasts, 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 elasticaFicus elastica
Ficus elastica, also called the rubber fig, rubber bush, rubber tree, rubber plant, or Indian rubber bush is a species of plant in the fig genus, native to northeast India and southern Indonesia....
and Peperomia
Peperomia
Peperomia is one of the 2 large genera of the Piperaceae family, with more than 1000 recorded species. Most of them are compact, small perennial epiphytes growing on rotten wood. More than 1500 species have been recorded, occurring in all tropical and subtropical regions of the world, though...
, 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
Cutin
Cutin is one of two waxy polymers that are the main components of the plant cuticle, which covers all aerial surfaces of plants. The other major cuticle polymer is cutan, which is much more readily preserved in the fossil record,...
, and are covered with a cuticle
Plant cuticle
Plant cuticles are a protective waxy covering produced only by the epidermal cells of leaves, young shoots and all other aerial plant organs without periderm...
. The cuticle reduces water loss to the atmosphere, it is sometimes covered with wax
Wax
thumb|right|[[Cetyl palmitate]], a typical wax ester.Wax refers to a class of chemical compounds that are plastic near ambient temperatures. Characteristically, they melt above 45 °C to give a low viscosity liquid. Waxes are insoluble in water but soluble in organic, nonpolar solvents...
in smooth sheets or long filaments. Thick 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. 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.
Trichome
Trichome
Trichomes are fine outgrowths or appendages on plants and certain protists. These are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae.- Algal trichomes :...
s 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
Secondary growth
In many vascular plants, secondary growth is the result of the activity of the two lateral meristems, the cork cambium and vascular cambium. Arising from lateral meristems, secondary growth increases the girth of the plant root or stem, rather than its length. As long as the lateral meristems...
, the epidermis of roots and stems is usually replaced by a periderm through the action of a cork cambium
Cork cambium
Cork cambium is a tissue found in many vascular plants as part of the periderm. The cork cambium is a lateral meristem and is responsible for secondary growth that replaces the epidermis in roots and stems...
.
Guard cells
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
Poaceae
The Poaceae is a large and nearly ubiquitous family of flowering plants. Members of this family are commonly called grasses, although the term "grass" is also applied to plants that are not in the Poaceae lineage, including the rushes and sedges...
, often have roughly equal numbers of stomata on both surfaces. The number of stomata varies from about 1,000 to over 100,000 per square centimeter of leaf surface.
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
The plant epidermis consists of three main cell types: pavement cells, guard cellGuard cell
Guard cells are specialized cells located in the Leaf epidermis of plants. Pairs of guard cells surround tiny stomatal airway pores . These tiny holes in the surface of leaves are necessary for gas exchange into and out of the plant; carbon dioxide enters the plant allowing the carbon fixation...
s and their subsidiary cells that surround the stomata and trichome
Trichome
Trichomes are fine outgrowths or appendages on plants and certain protists. These are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae.- Algal trichomes :...
s, 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
Genetics
Genetics , a discipline of biology, is the science of genes, heredity, and variation in living organisms....
and environmental conditions.
Trichomes develop at a distinct phase during the actual leaf
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
development, under the control of two major trichome specification gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...
s: TTG and GL1. The process may be controlled by the plant hormone
Plant hormone
Plant hormones are chemicals that regulate plant growth, which, in the UK, are termed 'plant growth substances'. Plant hormones are signal molecules produced within the plant, and occur in extremely low concentrations. Hormones regulate cellular processes in targeted cells locally and, when moved...
s gibberellin
Gibberellin
Gibberellins are plant hormones that regulate growth and influence various developmental processes, including stem elongation, germination, dormancy, flowering, sex expression, enzyme induction, and leaf and fruit senescence....
s, 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
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
without subsequent cell division
Cell division
Cell division is the process by which a parent cell divides into two or more daughter cells . Cell division is usually a small segment of a larger cell cycle. This type of cell division in eukaryotes is known as mitosis, and leaves the daughter cell capable of dividing again. The corresponding sort...
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
Arabidopsis thaliana
Arabidopsis thaliana is a small flowering plant native to Europe, Asia, and northwestern Africa. A spring annual with a relatively short life cycle, arabidopsis is popular as a model organism in plant biology and genetics...
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 petal
Petal
Petals are modified leaves that surround the reproductive parts of flowers. They often are brightly colored or unusually shaped to attract pollinators. Together, all of the petals of a flower are called a corolla. Petals are usually accompanied by another set of special leaves called sepals lying...
s of flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...
s. Expression of the gene MIXTA, or its analogue
Analog (chemistry)
In chemistry, a structural analog , also known as chemical analog or simply analog, is a compound having a structure similar to that of another one, but differing from it in respect of a certain component. It can differ in one or more atoms, functional groups, or substructures, which are replaced...
in other species, later in the process of cellular differentiation
Cellular differentiation
In developmental biology, cellular differentiation is the process by which a less specialized cell becomes a more specialized cell type. Differentiation occurs numerous times during the development of a multicellular organism as the organism changes from a simple zygote to a complex system of...
will cause the formation of conical cells over trichomes. MIXTA is a transcription factor
Transcription factor
In molecular biology and genetics, a transcription factor is a protein that binds to specific DNA sequences, thereby controlling the flow of genetic information from DNA to mRNA...
.
Stomatal patterning is a lot 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 mutant
Mutant
In biology and especially genetics, a mutant is an individual, organism, or new genetic character, arising or resulting from an instance of mutation, which is a base-pair sequence change within the DNA of a gene or chromosome of an organism resulting in the creation of a new character or trait not...
s do show it is under some form of genetic control, but it is more controlled in monocots, where stomata arise from specific asymmetric divisions
Asymmetric cell division
An asymmetric cell division produces two daughter cells with different cellular fates. This is in contrast to normal, symmetric, cell divisions, which give rise to daughter cells of equivalent fates...
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 and FLP 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
Density
The mass density or density of a material is defined as its mass per unit volume. The symbol most often used for density is ρ . In some cases , density is also defined as its weight per unit volume; although, this quantity is more properly called specific weight...
on the leaf surface. It is thought that plant hormones, such as ethylene
Ethylene
Ethylene is a gaseous organic compound with the formula . It is the simplest alkene . Because it contains a carbon-carbon double bond, ethylene is classified as an unsaturated hydrocarbon. Ethylene is widely used in industry and is also a plant hormone...
and cytokines, control the stomata’s 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
Plasmodesmata
Plasmodesmata are microscopic channels which traverse the cell walls of plant cells and some algal cells, enabling transport and communication between them. Species that have plasmodesmata include members of the Charophyceae, Charales and Coleochaetales , as well as all embryophytes, better known...
.
See also
- BarkBarkBark is the outermost layers of stems and roots of woody plants. Plants with bark include trees, woody vines and shrubs. Bark refers to all the tissues outside of the vascular cambium and is a nontechnical term. It overlays the wood and consists of the inner bark and the outer bark. The inner...
- Cork cambiumCork cambiumCork cambium is a tissue found in many vascular plants as part of the periderm. The cork cambium is a lateral meristem and is responsible for secondary growth that replaces the epidermis in roots and stems...
- Periderm