Leaf

A leaf (pl.: leaves) is a principal appendage of the stem of a vascular plant,^[1] usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage",^[2]^[3] while the leaves, stem, flower, and fruit collectively form the shoot system.^[4] In most leaves, the primary photosynthetic tissue is the palisade mesophyll and is located on the upper side of the blade or lamina of the leaf^[1] but in some species, including the mature foliage of Eucalyptus,^[5] palisade mesophyll is present on both sides and the leaves are said to be isobilateral. Most leaves are flattened and have distinct upper (adaxial) and lower (abaxial) surfaces that differ in color, hairiness, the number of stomata (pores that intake and output gases), the amount and structure of epicuticular wax and other features. Leaves are mostly green in color due to the presence of a compound called chlorophyll which is essential for photosynthesis as it absorbs light energy from the sun. A leaf with lighter-colored or white patches or edges is called a variegated leaf.

Leaves can have many different shapes, sizes, textures and colors. The broad, flat leaves with complex venation of flowering plants are known as megaphylls and the species that bear them, the majority, as broad-leaved or megaphyllous plants, which also include acrogymnosperms and ferns. In the lycopods, with different evolutionary origins, the leaves are simple (with only a single vein) and are known as microphylls.^[6] Some leaves, such as bulb scales, are not above ground. In many aquatic species, the leaves are submerged in water. Succulent plants often have thick juicy leaves, but some leaves are without major photosynthetic function and may be dead at maturity, as in some cataphylls and spines. Furthermore, several kinds of leaf-like structures found in vascular plants are not totally homologous with them. Examples include flattened plant stems called phylloclades and cladodes, and flattened leaf stems called phyllodes which differ from leaves both in their structure and origin.^[3]^[7] Some structures of non-vascular plants look and function much like leaves. Examples include the phyllids of mosses and liverworts.

General characteristics

3D rendering of a computed tomography scan of a leaf

Leaves are the most important organs of most vascular plants.^[8] Green plants are autotrophic, meaning that they do not obtain food from other living things but instead create their own food by photosynthesis. They capture the energy in sunlight and use it to make simple sugars, such as glucose and sucrose, from carbon dioxide and water. The sugars are then stored as starch, further processed by chemical synthesis into more complex organic molecules such as proteins or cellulose, the basic structural material in plant cell walls, or metabolized by cellular respiration to provide chemical energy to run cellular processes. The leaves draw water from the ground in the transpiration stream through a vascular conducting system known as xylem and obtain carbon dioxide from the atmosphere by diffusion through openings called stomata in the outer covering layer of the leaf (epidermis), while leaves are orientated to maximize their exposure to sunlight. Once sugar has been synthesized, it needs to be transported to areas of active growth such as the plant shoots and roots. Vascular plants transport sucrose in a special tissue called the phloem. The phloem and xylem are parallel to each other, but the transport of materials is usually in opposite directions. Within the leaf these vascular systems branch (ramify) to form veins which supply as much of the leaf as possible, ensuring that cells carrying out photosynthesis are close to the transportation system.^[9]

Typically leaves are broad, flat and thin (dorsiventrally flattened), thereby maximising the surface area directly exposed to light and enabling the light to penetrate the tissues and reach the chloroplasts, thus promoting photosynthesis. They are arranged on the plant so as to expose their surfaces to light as efficiently as possible without shading each other, but there are many exceptions and complications. For instance, plants adapted to windy conditions may have pendent leaves, such as in many willows and eucalypts. The flat, or laminar, shape also maximizes thermal contact with the surrounding air, promoting cooling. Functionally, in addition to carrying out photosynthesis, the leaf is the principal site of transpiration, providing the energy required to draw the transpiration stream up from the roots, and guttation.

Many conifers have thin needle-like or scale-like leaves that can be advantageous in cold climates with frequent snow and frost.^[10] These are interpreted as reduced from megaphyllous leaves of their Devonian ancestors.^[6] Some leaf forms are adapted to modulate the amount of light they absorb to avoid or mitigate excessive heat, ultraviolet damage, or desiccation, or to sacrifice light-absorption efficiency in favor of protection from herbivory. For xerophytes the major constraint is not light flux or intensity, but drought.^[11] Some window plants such as Fenestraria species and some Haworthia species such as Haworthia tesselata and Haworthia truncata are examples of xerophytes.^[12] and Bulbine mesembryanthemoides.^[13]

Leaves also function to store chemical energy and water (especially in succulents) and may become specialized organs serving other functions, such as tendrils of peas and other legumes, the protective spines of cacti and the insect traps in carnivorous plants such as Nepenthes and Sarracenia.^[14] Leaves are the fundamental structural units from which cones are constructed in gymnosperms (each cone scale is a modified megaphyll leaf known as a sporophyll)^[6]^: 408 and from which flowers are constructed in flowering plants.^[6]^: 445

The internal organization of most kinds of leaves has evolved to maximize exposure of the photosynthetic organelles, the chloroplasts, to light and to increase the absorption of carbon dioxide while at the same time controlling water loss. Their surfaces are waterproofed by the plant cuticle and gas exchange between the mesophyll cells and the atmosphere is controlled by minute (length and width measured in tens of μm) openings called stomata which open or close to regulate the rate exchange of carbon dioxide(CO₂), oxygen(O₂) and water vapor into and out of the internal intercellular space system. Stomatal opening is controlled by the turgor pressure in a pair of guard cells that surround the stomatal aperture. In any square centimeter of a plant leaf, there may be from 1,000 to 100,000 stomata.^[15]

The shape and structure of leaves vary considerably from species to species of plant, depending largely on their adaptation to climate and available light, but also to other factors such as grazing animals (such as deer), available nutrients, and ecological competition from other plants. Considerable changes in leaf type occur within species, too, for example as a plant matures; as a case in point Eucalyptus species commonly have isobilateral, pendent leaves when mature and dominating their neighbors; however, such trees tend to have erect or horizontal dorsiventral leaves as seedlings, when their growth is limited by the available light.^[16] Other factors include the need to balance water loss at high temperature and low humidity against the need to absorb atmospheric carbon dioxide. In most plants, leaves also are the primary organs responsible for transpiration and guttation (beads of fluid forming at leaf margins).

Leaves can also store food and water, and are modified accordingly to meet these functions, for example in the leaves of succulent plants and in bulb scales. The concentration of photosynthetic structures in leaves requires that they be richer in protein, minerals, and sugars than, say, woody stem tissues. Accordingly, leaves are prominent in the diet of many animals.

Correspondingly, leaves represent heavy investment on the part of the plants bearing them, and their retention or disposition are the subject of elaborate strategies for dealing with pest pressures, seasonal conditions, and protective measures such as the growth of thorns and the production of phytoliths, lignins, tannins and poisons.

Deciduous plants in frigid or cold temperate regions typically shed their leaves in autumn, whereas in areas with a severe dry season, some plants may shed their leaves until the dry season ends. In either case, the shed leaves may be expected to contribute their retained nutrients to the soil where they fall.

In contrast, many other non-seasonal plants, such as palms and conifers, retain their leaves for long periods; Welwitschia retains its two main leaves throughout a lifetime that may exceed a thousand years.

The leaf-like organs of bryophytes (e.g., mosses and liverworts), known as phyllids, differ heavily morphologically from the leaves of vascular plants. In most cases, they lack vascular tissue, are only a single cell thick, and have no cuticle, stomata, or internal system of intercellular spaces. (The phyllids of the moss family Polytrichaceae are notable exceptions.) The phyllids of bryophytes are only present on the gametophytes, while in contrast the leaves of vascular plants are only present on the sporophytes. These can further develop into either vegetative or reproductive structures.^[14]

Simple, vascularized leaves (microphylls), such as those of the early Devonian lycopsid Baragwanathia, first evolved as enations, extensions of the stem. True leaves or euphylls of larger size and with more complex venation did not become widespread in other groups until the Devonian period, by which time the carbon dioxide concentration in the atmosphere had dropped significantly. This occurred independently in several separate lineages of vascular plants, in progymnosperms like Archaeopteris, in Sphenopsida, ferns and later in the gymnosperms and angiosperms. Euphylls are also referred to as macrophylls or megaphylls (large leaves).^[6]

Morphology

Animated zoom into the leaf of a Sequoia sempervirens (California redwood)

A structurally complete leaf of an angiosperm consists of a petiole (leaf stalk), a lamina (leaf blade), stipules (small structures located to either side of the base of the petiole) and a sheath. Not every species produces leaves with all of these structural components. The proximal stalk or petiole is called a stipe in ferns. The lamina is the expanded, flat component of the leaf which contains the chloroplasts. The sheath is a structure, typically at the base that fully or partially clasps the stem above the node, where the leaf is attached. Leaf sheathes typically occur in Poaceae (grasses) and Apiaceae (umbellifers). Between the sheath and the lamina, there may be a pseudopetiole, a petiole like structure. Pseudopetioles occur in some monocotyledons including bananas, palms and bamboos.^[18] Stipules may be conspicuous (e.g. beans and roses), soon falling or otherwise not obvious as in Moraceae or absent altogether as in the Magnoliaceae. A petiole may be absent (apetiolate), or the blade may not be laminar (flattened). The petiole mechanically links the leaf to the plant and provides the route for transfer of water and sugars to and from the leaf. The lamina is typically the location of the majority of photosynthesis. The upper (adaxial) angle between a leaf and a stem is known as the axil of the leaf. It is often the location of a bud. Structures located there are called "axillary".

External leaf characteristics, such as shape, margin, hairs, the petiole, and the presence of stipules and glands, are frequently important for identifying plants to family, genus or species levels, and botanists have developed a rich terminology for describing leaf characteristics. Leaves almost always have determinate growth. They grow to a specific pattern and shape and then stop. Other plant parts like stems or roots have non-determinate growth, and will usually continue to grow as long as they have the resources to do so.

The type of leaf is usually characteristic of a species (monomorphic), although some species produce more than one type of leaf (dimorphic or polymorphic). The longest leaves are those of the Raffia palm, R. regalis which