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Look up foliage in Wiktionary, the free dictionary.
In botany, a leaf is an above-ground plant organ specialized for photosynthesis. For this purpose, a leaf is typically flat (laminar) and thin, to expose the cells containing chloroplast (chlorenchyma tissue, a type of parenchyma) to light over a broad area, and to allow light to penetrate fully into the tissues. Leaves are also the sites in most plants where transpiration and guttation take place. Leaves can store food and water, and are modified in some plants for other purposes. The comparable structures of ferns are correctly referred to as fronds. Furthermore, leaves are prominent in the human diet as leaf vegetables.
Additional recommended knowledge
A structurally complete leaf of an angiosperm consists of a petiole (leaf stem), a lamina (leaf blade), and stipules (small processes located to either side of the base of the petiole). The petiole attaches to the stem at a point called the "leaf axil". Not every species produces leaves with all of the afformentioned structural components. In some species, paired stipules are not obvious or are absent altogether. A petiole may be absent, or the blade may not be laminar (flattened). The tremendous variety shown in leaf structure (anatomy) from species to species is presented in detail below under Leaf morphology. After a period of time (i.e. seasonally, during the autumn), deciduous trees shed their leaves. These leaves then decompose into the soil.
A leaf is considered a plant organ and typically consists of the following tissues:
The epidermis is the outer multi-layered group of cells covering the leaf. It forms the boundary separating the plant's inner cells from the external world. The epidermis serves several functions: protection against water loss, regulation of gas exchange, secretion of metabolic compounds, and (in some species) absorption of water. Most leaves show dorsoventral anatomy: the upper (adaxial) and lower (abaxial) surfaces have somewhat different construction and may serve different functions.
The epidermis is usually transparent (epidermal cells lack chloroplasts) and coated on the outer side with a waxy cuticle that prevents water loss. The cuticle is in some cases thinner on the lower epidermis than on the upper epidermis, and is thicker on leaves from dry climates as compared with those from wet climates.
The epidermis 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.
The epidermis is covered with pores called stomata, 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 than the adaxial (upper) epidermis.
Most of the interior of the leaf between the upper and lower layers of epidermis is a parenchyma (ground tissue) or chlorenchyma tissue called the mesophyll (Greek for "middle leaf"). This assimilation tissue is the primary location of photosynthesis in the plant. The products of photosynthesis are called "assimilates".
In ferns and most flowering plants the mesophyll is divided into two layers:
The pores or stomata of the epidermis open into substomatal chambers, connecting to air spaces between the spongy layer cells.
These two different layers of the mesophyll are absent in many aquatic and marsh plants. Even an epidermis and a mesophyll may be lacking. Instead for their gaseous exchanges they use a homogeneous aerenchyma (thin-walled cells separated by large gas-filled spaces). Their stomata are situated at the upper surface.
Leaves in temperate, boreal, and seasonally dry zones may be seasonally deciduous (falling off or dying for the inclement season). This mechanism to shed leaves is called abscission. After the leaf is shed, a leaf scar develops on the twig. In cold autumns they sometimes change color, and turn yellow, bright orange or red as various accessory pigments (carotenoids and xanthophylls) are revealed when the tree responds to cold and reduced sunlight by curtailing chlorophyll production. Red anthocyanin pigments are now thought to be produced in the leaf as it dies.
The veins are the vascular tissue of the leaf and are located in the spongy layer of the mesophyll. They are typical examples of pattern formation through ramification. The pattern of the veins is called venation.
The veins are made up of:
The xylem typically lies over the phloem. Both are embedded in a dense parenchyma tissue, called "pith", with usually some structural collenchyma tissue present.
External leaf characteristics (such as shape, margin, hairs, etc.) are important for identifying plant species, and botanists have developed a rich terminology for describing leaf characteristics. These structures are a part of what makes leaves determinant, they grow and achieve a specific pattern and shape, then stop. Other plant parts like stems or roots are non-determinant, and will usually continue to grow as long as they have the resources to do so.
Classification of leaves can occur through many different designative schema, and the type of leaf is usually characteristic of a species, although some species produce more than one type of leaf. The longest type of leaf is a leaf from palm trees, measuring at nine feet long. The terminology associated with the description of leaf morphology is presented, in illustrated form, at Wikibooks.
Basic leaf types
Arrangement on the stem
Different terms are usually used to describe leaf placement (phyllotaxis):
As a stem grows, leaves tend to appear arranged around the stem in a way that optimizes yield of light. In essence, leaves form a helix pattern centred around the stem, either clockwise or counterclockwise, with (depending upon the species) the same angle of divergence. There is a regularity in these angles and they follow the numbers in a Fibonacci sequence: 1/2, 2/3, 3/5, 5/8, 8/13, 13/21, 21/34, 34/55, 55/89. This series tends to a limit of 360° x 34/89 = 137.52 or 137° 30', an angle known mathematically as the golden angle. In the series, the numerator indicates the number of complete turns or "gyres" until a leaf arrives at the initial position. The denominator indicates the number of leaves in the arrangement. This can be demonstrated by the following:
The fact that an arrangement of anything in nature can be described by a mathematical formula is not in itself mysterious. Mathematics are the science of discovering numerical relationships and applying formulae to these relationships. The formulae themselves can provide clues to the underlying physiological processes that, in this case, determine where the next leaf bud will form in the elongating stem.
Divisions of the lamina (blade)
Two basic forms of leaves can be described considering the way the blade is divided. A simple leaf has an undivided blade. However, the leaf shape may be formed of lobes, but the gaps between lobes do not reach to the main vein. A compound leaf has a fully subdivided blade, each leaflet of the blade separated along a main or secondary vein. Because each leaflet can appear to be a simple leaf, it is important to recognize where the petiole occurs to identify a compound leaf. Compound leaves are a characteristic of some families of higher plants, such as the Fabaceae. The middle vein of a compound leaf or a frond, when it is present, is called a rachis.
Characteristics of the petiole
Petiolated leaves have a petiole. Sessile leaves do not: the blade attaches directly to the stem. In clasping or decurrent leaves, the blade partially or wholly surrounds the stem, often giving the impression that the shoot grows through the leaf. When this is actually the case, the leaves are called "perfoliate", such as in Claytonia perfoliata. In peltate leaves, the petiole attaches to the blade inside from the blade margin.
In some Acacia species, such as the Koa Tree (Acacia koa), the petioles are expanded or broadened and function like leaf blades; these are called phyllodes. There may or may not be normal pinnate leaves at the tip of the phyllode.
A stipule, present on the leaves of many dicotyledons, is an appendage on each side at the base of the petiole resembling a small leaf. Stipules may be lasting and not be shed (a stipulate leaf, such as in roses and beans), or be shed as the leaf expands, leaving a stipule scar on the twig (an exstipulate leaf).
Venation (arrangement of the veins)
There are two subtypes of venation, namely, craspedodromous, where the major veins stretch up to the margin of the leaf, and camptodromous, when major veins extend close to the margin, but bend before they intersect with the margin.
Note that although it is the more complex pattern, branching veins appear to be plesiomorphic and in some form were present in ancient seed plants as long as 250 million years ago. A pseudo-reticulate venation that is actually a highly modified penniparallel one is an autapomorphy of some Melanthiaceae which are monocots, e.g. Paris quadrifolia (True-lover's Knot).
Leaf morphology changes within a single plant
The leaf margin is characteristic for a genus and aids in determining the species.
Tip of the leaf
Base of the leaf
Surface of the leaf
The surface of a leaf can be described by several botanical terms:
The leaf surface is also host to a large variety of microorganisms; in this context it is referred to as the phyllosphere.
"Hairs" on plants are properly called trichomes. Leaves can show several degrees of hairiness. The meaning of several of the following terms can overlap.
In the course of evolution, leaves adapted to different environments in the following ways:
Interactions with other organisms
Although not as nutritious as other organs such as fruit, leaves provide a food source for many organisms. Animals which eat leaves are known as folivores. The leaf is one of the most vital parts of the plant, and plants have evolved protection against folivores such as tannins, chemicals which hinder the digestion of proteins and have an unpleasant taste. Some animals have cryptic adaptations to avoid their own predators, for example some caterpillars will create a small home in the leaf by folding it over themselves, while other herbivores and their prey mimic the appearance of the leaf. Some insects, such as the katydid, take this even further, moving from side to side much like a leaf does in the wind.
|This article is licensed under the GNU Free Documentation License. It uses material from the Wikipedia article "Leaf". A list of authors is available in Wikipedia.|