This article is about the leaf, a plant organ. See Leaf (disambiguation) for other meanings.

Fallen leaf of a maple. Note areas where chlorophyll (green) has been destroyed now appear yellow.

In botany, a leaf is an above-ground plant organ specialized for photosynthesis. For this purpose, a leaf is typically flat and thin, to expose the chloroplast containing cells (chlorenchyma) 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 respiration, 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 referred to as fronds.

Leaf structure

Detailed view of the pinnate veins

A structurally complete leaf of an angiosperm consists typically of a petiole (leaf stem), a lamina (leaf blade), and stipules (small processes located to either side of the base of the petiole).

The point at which the petiole attaches to the plant stem is called the leaf axil.

Not all species produce leaves with all of these parts. In some species, stipules are not obvious or present, or the petiole may be absent. The blade is not always laminar (flattened).

A leaf has a dorsiventral anatomy, in other words the upper surface and the underside have a different construction and different functions.

The external leaf characteristics (such as shape, margin, hairs, etc.) are important for identifying plant species.

A leaf typically consists of the following tissues:

1. An epidermis that covers the upper and lower surfaces
2. An interior chlorenchyma called the mesophyll
3. A number of veins (the vascular tissue).

Epidermis

The epidermis is the outer multi-layered group of cells covering the leaf blade. It is the boundary between the plant and its surroundings. The structure of the epidermis of the upper surface and the underside of the leaf may be different.

The epidermis has several functions: absorption of water, protection against water loss, gas exchange through photosynthesis, secretion of metabolic compounds, and protection against outside influences.

The layer is usually transparent (cells lack chloroplasts) and coated on the outer surface with a waxy cuticle that prevents water loss. The cuticle may be 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 contains differentiated cell types; epidermal cells, guard cells, subsidiary cells and the epidermal hairs (trichomes).

The epidermal cells are the largest and the least specialized group. They are elongated in the leaves of monocots.

The epidermis is covered with pores called stomata (sing., stoma). It is part of the stoma complex: the stoma, enclosed at each side by chloroplast-containing guard cells, and two to four subsidiary cells (lacking chloroplast). These stomata enable oxygen and carbon dioxide to move in and out of the leaf. There is a substomatal chamber (an air space) below the stoma. These pores are more numerous over the abaxial (lower) epidermis than the (adaxial) upper epidermis in most leaves. Water vapour also passes out of the stomata during transpiration.

To conserve water, the paired guard cells at each side of the stomata may force the opening or the closing of the stomata. This happens through a differential thickening of the walls of the guard cells. When they become inflated in a humid surrounding, the internal water pressure (turgor pressure) opens up the stoma. The inverse happens, as the guard cells lose water pressure on a warm day, regulating transpiration and the flow of carbon dioxide.

The opening and closing of the stomata is also triggered by light. They are open during daytime and closed at night. The reverse happens with some plants from arid regions, such as some of the sedum family (Crassulaceae); to prevent extensive water loss, they open their stomata during the night.

Stomata are not found in some underwater plants.

Trichomes or hairs grow out from the epidermis in many species.

Mesophyll

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 (= middle leaf). This assimilation tissue is made of chloroplast and is the primary photosynthetic tissue of the plant. The products of this photosynthesis are called assimilates. There are also a few plants with mesophyll which lack chloroplasts, and therefore also lack assimilation.

In ferns and most flowering plants it is divided into two layers:

• an upper palisade layer of tightly packed, vertically elongated cells, one to two cells thick, directly beneath the adaxial epidermis. Its cells contain many more chloroplasts than the spongy layer. These long cylindrical cells are regularly arranged in one to five rows. Cylindrical cells, with the chloroplasts close to the walls of the cell, can take optimal advantage of light. The slight separation of the cells provides maximal absorption of carbon dioxide. This separation must be minimal to afford capillary action for water distribution. In order to adapt to their different environment (such as sun or shade), plants had to adapt this structure to obtain optimal result. Sun leaves have a multi-layered palisade layer, while shade leaves or older leaves closer to the soil, are single-layered.
• Beneath the palisade layer is the spongy layer. The cells of the spongy layer are more rounded and not so tightly packed. There are large intercellular air spaces (substomatal chambers). These cells contain less chloroplasts.

The pores or stomata of the epidermis open into substomatal chambers, the 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 homogenous aerenchyma (thin-walled cells separated by large gas-filled spaces). Their stomata are situated at the upper surface.

Leaves are normally green in color, which comes from chlorophyll found in plastids in the chlorenchyma.

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 turn yellow, bright orange or red as various accessory pigments (carotenoids and anthocyanins) are revealed when the tree responds to cold and reduced sunlight by curtailing chlorophyll production.

Veins

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 veins are made up of:

• xylem, which brings water from the stem into the leaf.
• phloem, which usually moves sap out, the latter containing the glucose produced by photosynthesis in the leaf.

The xylem typically lies over the phloem. Both are embedded in a dense parenchyma tissue (= ground tissue), called pith, with usually some structural collenchyma tissue present.

Leaf Types, Arrangements, and Forms

The leaves on this plant are arranged in pairs opposite one another, with successive pairs at right angles to each other ("decussate") along the red stem. Note developing buds in the axils of these leaves.

Leaves may be classified in many different ways, and the type is usually characteristic of a species, although some species produce more than one type of leaf. The terminology associated with describing leaf morphology is presented (with illustrations) at Wikibooks (http://wikibooks.org/wiki/Botany:_Leaves_(forms)).

• Basic leaf types:
  • Ferns have fronds.
  • Conifer leaves are typically needle-, awl-, or scale-shaped
  • Angiosperm (flowering plant) leaves: the standard form includes stipules, petiole, and lamina.
  • Microphyll leaves.
  • Sheath leaves (type found in most grasses).
  • Specialized leaves.

• Arrangement on the stem
  • Spiral — leaf attachments arranged in interlocking spirals, with the spirals in Fibonacci number ratios (1:1:2:3:5:8:13:21:etc; e.g. five successive leaves counted along a shoot clockwise, and eight leaves anti-clockwise, come to the same leaf).
  • Alternate — leaf attachments singular at nodes, and alternate direction up the stem.
  • Opposite — leaf attachments paired at each node; decussate if, as typical, each successive pair is rotated 90� going along the stem; or distichous if not rotated, but all two-ranked, in the same plane.
  • Whorled — three or more leaves attach at each point or node on the stem. As with opposite leaves, successive whorls may or may not be decussate, rotated by half the angle between the leaves in the whorl (i.e., successive whorls of three rotated 60�, whorls of four rotated 45�, etc). Note: opposite leaves may appear whorled near the tip of the stem.
  • Rosulate — leaves form a rosette ( = a cluster of leaves growing in crowded circles from a common center).

Leaves of the Norway Spruce (Picea abies) are needle-shaped and the arrangement is spiral

Leaves are set in a particular arrangement around the stem in order to gain an optimal yield of light. Subsequent leaves are arranged in spirals, clockwise or counterclockwise, with always the same angle of divergence. There is a certain regularity in these angles: they follow the numbers in a Fibonacci series: 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'. This angle is known mathematically as the 'golden angle'. The numerator gives the number of gyres, till the leaf arrives at the same initial position. The denominator gives the number of leaves in this arrangement. This can easily be seen.

• Divisions of the lamina (blade):
  • Simple leaves have an undivided blade. The leaf shape may be one of deeply divided lobes, but the gaps between lobes do not reach to the vein.
  • Compound leaves have divided blades, each leaflet separated along a main or secondary vein.
    • Palmately compound leaves have the leaflets radiating from the end of the petiole, like fingers off the palm of a hand. There is no rachis, e.g. Cannabis (hemp) and Aesculus (buckeyes).
    • Pinnately compound leaves have the leaflets arranged along the main or mid-vein (called a rachis in this case).
      • odd pinnate: with a terminal leaflet, e.g. Fraxinus (ash).
      • even pinnate: lacking a terminal leaflet, e.g. Swietenia (mahogany).
    • Bipinnately compound leaves are twice divided: the leaflets are arranged along a secondary vein that is one of several branching off the rachis. Each leaflet is called a pinnule. The pinnules on one secondary vein are called pinna; e.g. Albizia (silk tree).
    • trifoliate: a pinnate leaf with just three leaflets, e.g. Trifolium (clover), Laburnum (laburnum).
    • pinnatifid: pinnately dissected to the midrib, but with the leaflets not entirely separate, e.g. some Sorbus (whitebeams).

• Characteristics of the petiole:
  • Petiolated leaves have a petiole.
    • In peltate leaves, the petiole attaches to the blade inside from the blade margin.
  • Sessile or clasping leaves do not have a petiole. In sessile leaves the blade attaches directly to the stem. In clasping leaves, the blade partially or wholly surrounds the stem, giving the impression that the shoot grows through the leaf such as in Claytonia perfoliata of the purslane family (Portulacaceae).

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.

• Characteristics of the stipule
  • 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. They 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).
  • The situation, arrangement, and structure of the stipules is called the stipulation.
    • free
    • adnate : fused to the petiole base
    • ochreate : provided with ochrea, or sheath-formed stipules, e.g. rhubarb,
    • encircling the petiole base
    • interpetiolar : between the petioles of two opposite leaves.
    • intrapetiolar : between the petiole and the subtending stem

• Arrangement of the veins (venation):

There are two subtypes of venation, craspedodromus (the major veins stretch up to the margin of the leaf) and camptodromous (major veins come close to the margin, but bend before they get to it).

• • Feather-veined, reticulate — the veins arise pinnately from a single mid-vein and subdivide into veinlets. These, in turn, form a complicated network. This type of venation is typical for dicotyledons.
    • Pinnate-netted, penniribbed, penninerved, penniveined; the leaf has usually one main vein (called the mid-vein), with veinlets, smaller veins branching off laterally, usually somewhat parallel to each other; eg Malus (apples).
    • Three main veins originate from the base of the lamina, as in Ceanothus.
    • Palmate-netted, palmate-veined, fan-veined; several main veins diverge from near the leaf base where the petiole attaches, and radiate toward the edge of the leaf; e.g. most Acer (maples).
  • Parallel-veined, parallel-ribbed, parallel-nerved, penniparallel — veins run parallel most the length of the leaf, from the base to the apex. Commissural veins (small veins) connect the major parallel veins. Typical for most monocotyledons, such as grasses.
  • Dichotomous — There are no dominant bundles, with the veins forking regularly by pairs; found in Ginkgo and some pteridophytes.

Underside view of leaf

Leaf shapes

See Leaf shape

Leaf margins

The leaf margin is characteristic for a genus and aids in determining the species.

• entire: even; with a smooth margin; without toothing
• ciliate: fringed with hairs
• crenate: wavy-toothed; dentate with rounded teeth, such as Fagus (beech)
• dentate: toothed, such as Castanea (chestnut)
  • coarse-toothed: with large teeth
  • glandular toothed: with teeth that bear glands.
• denticulate: finely toothed
• doubly toothed: each tooth bearing smaller teeth, such as Ulmus (elm)
• lobate: indented, with the indentations not reaching to the center, such as many Quercus (oaks)
  • palmately lobed: indented with the indentations reaching to the center, such as Humulus (hop).
• serrate: saw-toothed with asymmetrical teeth pointing forward, such as Urtica (nettle)
• serrulate: finely serrate
• sinuate: with deep, wave-like indentations; coarsely crenate, such as many Rumex (docks)
• spiny: with stiff, sharp points, such as some Ilex (hollies) and Cirsium (thistles).

Leaf tips

• acuminate: long-pointed, prolonged into a narrow, tapering point in a concave manner.
• acute: ending in a sharp, but not prolonged point
• cuspidate: with a sharp, elongated, rigid tip; tipped with a cusp.
• emarginate: indented, with a shallow notch at the tip.
• mucronate: abruptly tipped with a small short point, as a continuation of the midrib; tipped with a mucro.
• mucronulate: mucronate, but with a smaller spine.
• obcordate: inversely heart-shaped, deeply notched at the top.
• obtuse: rounded or blunt
• truncate: ending abruptly with a flat end, that looks cut off.

Leaf bases

• acuminate: coming to a sharp, narrow, prolonged point.
• acute: coming to a sharp, but not prolonged point.
• auriculate: ear-shaped
• cordate: heart-shaped with the norch away from the stem.
• cuneate: wedge-shaped.
• hastate: shaped like an halberd and with the basal lobes pointing outward.
• oblique: slanting.
• reniform: kidney-shaped but rounder and broader than long.
• rounded: curving.
• sagittate: shaped like an arrowhead and with the acute basal lobes pointing downward.
• truncate: ending abruptly with a flat end, that looks cut off.

Hairiness

Leaves can show several degrees of hairiness. The meaning of several of the following terms can overlap.

• glabrous: no hairs of any kind present.
• arachnoid, arachnose: with many fine, entangled hairs giving a cobwebby appearance.
• barbellate: with finely barbed hairs (barbellae).
• bearded: with long, stiff hairs.
• bristly: with stiff hair-like prickles.
• canescent: hoary with dense grayish-white pubescence.
• ciliate: marginally fringed with short hairs (cilia).
• ciliolate: minutely ciliate.
• floccose: with flocks of soft, woolly hairs, which tend to rub off.
• glandular: with a gland at the tip of the hair.
• hirsute: with rather rough or stiff hairs.
• hispid: with rigid, bristly hairs.
• hispidulous: minutely hispid.
• hoary: with a fine, close grayish-white pubescence.
• lanate, lanose: with woolly hairs.
• pilose: with soft, clearly separated hairs.
• puberulent, puberulous: with fine, minute hairs.
• pubescent: with soft, short and erect hairs.
• scabrous, scabrid: rough to the touch
• sericeous: silky appearance through fine, straight and appressed (lying close and flat) hairs.
• silky: with adpressed, soft and straight pubescence.
• stellate, stellifrom: with star-shaped hairs.
• strigose: with appressed, sharp, straight and stiff hairs.
• tomentose: densely pubescent with matted, soft white woolly hairs.
  • cano-tomentose: between canescent and tomentose
  • felted-tomentose: woolly and matted with curly hairs.
• villous: with long and soft hairs, usually curved.
• woolly: with long, soft and tortuous or matted hairs.

Leaf surfaces

The surface of a leaf can be described by several botanical terms:

• farinose: bearing farina; mealy, covered with a waxy, whitish powder.
• glabrous: smooth, not hairy.
• glaucous: with a whitish bloom; covered with a very fine, bluish-white powder.
• glutinous: sticky, viscid.
• papillate, papillose: bearing papillae (minute, nipple-shaped protuberances).
• pubescent: covered with erect hairs (especially soft and short ones)
• punctate: marked with dots; dotted with depressions or with translucent glands or colored dots.
• rugose: deeply wrinkled; with veins clearly visible.
• scurfy: covered with tiny, broad scalelike particles.
• tuberculate: covered with tubercles; covered with warty prominences.
• verrucose: warted, with warty outgrowths.
• viscid, viscous: covered with thick, sticky secretions.

Adaptations

In order to survive in a harsh environment, leaves can adapt in the following ways:

• Hairy leaf surface to lessen water loss
• Waxy leaf surface to prevent water loss
• Small, shiny leaves to deflect the sun's rays
• Thicker leaves to store water (e.g. rhubarb)
• Spines instead of leaves (e.g. cactus)
• Leaves to trap insects (e.g. pitcher plant)
• Bulbs to store food (e.g. onion)

See Also

• Cuneate
• Leaf blower
• Vernation

External link

• Position and Arrangement (http://www.ibiblio.org/botnet/glossary/b_i.html)

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