Aleaf (pl.:leaves) is a principal appendage of thestem of avascular plant,[1] usually borne laterally above ground and specialized forphotosynthesis. Leaves are collectively calledfoliage, as in "autumn foliage",[2][3] while the leaves, stem,flower, andfruit collectively form theshoot system.[4] In most leaves, the primaryphotosynthetictissue is thepalisade 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 ofEucalyptus,[5] palisade mesophyll is present on both sides and the leaves are said to be isobilateral. The leaf is an integral part of the stem system, and most leaves are flattened and have distinct upper (adaxial) and lower (abaxial) surfaces that differ in color,hairiness, the number ofstomata (pores that intake and output gases), the amount and structure ofepicuticular wax, and other features. Leaves are mostly green in color due to the presence of a compound calledchlorophyll which is essential for photosynthesis as it absorbs light energy from theSun. A leaf with lighter-colored or white patches or edges is called avariegated leaf.
Leaves can have many different shapes, sizes, textures and colors. The broad, flat leaves with complexvenation offlowering plants are known asmegaphylls and the species that bear them (the majority) as broad-leaved ormegaphyllous plants, which also includeacrogymnosperms andferns. In thelycopods, with different evolutionary origins, the leaves are simple (with only a single vein) and are known asmicrophylls.[6] Some leaves, such asbulb 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 somecataphylls andspines. Furthermore, several kinds of leaf-like structures found in vascular plants are not totally homologous with them. Examples include flattened plant stems calledphylloclades andcladodes, and flattened leaf stems calledphyllodes 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 thephyllids ofmosses andliverworts.
Leaves are the most important organs of mostvascular plants.[8] Green plants areautotrophic, meaning that they do not obtain food from other living things but instead create their own food byphotosynthesis. They capture the energy insunlight and use it to make simplesugars, such asglucose andsucrose, fromcarbon dioxide (CO2) and water. The sugars are then stored asstarch, further processed bychemical synthesis into more complex organic molecules such asproteins orcellulose, the basic structural material in plant cell walls, ormetabolized bycellular respiration to provide chemical energy to run cellular processes. The leaves draw water from the ground in thetranspiration stream through avascular conducting system known asxylem and obtain carbon dioxide from theatmosphere by diffusion through openings calledstomata 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 theshoots androots. Vascular plants transport sucrose in a special tissue called thephloem. 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 thatcells 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 thetissues and reach thechloroplasts, 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 havependent leaves, such as in manywillows andeucalypts. The flat, or laminar, shape also maximizesthermal contact with the surrounding air, promoting cooling. Functionally, in addition to carrying out photosynthesis, the leaf is the principal site oftranspiration, providing the energy required to draw the transpiration stream up from the roots, andguttation.
Manyconifers 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 frommegaphyllous leaves of theirDevonian 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. Forxerophytes the major constraint is not lightflux orintensity, but drought.[11] Somewindow plants such asFenestraria species and someHaworthia species such asHaworthia tesselata andHaworthia truncata are examples of xerophytes.[12]
Leaves function to store chemical energy and water (especially insucculents) and may become specialized organs serving other functions, such astendrils ofpeas and otherlegumes, the protectivespines ofcacti, and the insect traps incarnivorous plants such asNepenthes andSarracenia.[13] Leaves are the fundamental structural units from whichcones are constructed ingymnosperms (each cone scale is a modified megaphyll leaf known as asporophyll)[6]: 408 and from which flowers are constructed inflowering plants.[6]: 445
Vein skeleton of a leaf. Veins containlignin that make them harder to degrade for microorganisms.
The internal organization of most kinds of leaves has evolved to maximize exposure of the photosyntheticorganelles (chloroplasts) to light and to increase the absorption of CO2 while at the same time controlling water loss. Their surfaces are waterproofed by theplant cuticle, and gas exchange between the mesophyll cells and the atmosphere is controlled by minute (length and width measured in tens of μm) stomata which open or close to regulate the rate exchange of CO2,oxygen (O2), andwater vapor into and out of the internal intercellular space system. Stomatal opening is controlled by theturgor pressure in a pair ofguard cells that surround the stomatal aperture. In any square centimeter of a plant leaf, there may be from 1,000 to 100,000 stomata.[14]
Near the ground theseEucalyptus saplings have juvenile dorsiventral foliage from the previous year, but this season their newly sprouting foliage is isobilateral, like the mature foliage on the adult trees above
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, available nutrients, and ecological competition from other plants. Considerable changes in leaf type occur within species, too, for example as a plant matures (Eucalyptus species commonly have isobilateral, pendent leaves when mature and dominating their neighbors; however, such trees tend to have erect or horizontaldorsiventral leaves as seedlings, when their growth is limited by the available light.)[15] Other factors include the need to balance water loss at high temperature and low humidity against the need to absorb CO2. In most plants, leaves also are the primary organs responsible fortranspiration andguttation (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 inbulb scales. The concentration of photosynthetic structures in leaves requires that they be richer inprotein,minerals, and sugars than, say, woody stem tissues. Accordingly, leaves are prominent in thediet of manyanimals. 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 ofphytoliths,lignins,tannins andpoisons.
Deciduous plants in cold temperate regions typically shed their leaves inautumn, whereas in areas with a severedry season, some plants may shed their leaves until the dry season ends. In either case, the shed leaves often contribute their retained nutrients to the soil where they fall. In contrast, many other non-seasonal plants, such aspalms 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 ofbryophytes (e.g.,mosses andliverworts), known asphyllids, differ greatly morphologically from the leaves ofvascular plants. In most cases, they lack vascular tissue, are a single cell thick and have nocuticle, stomata, or internal system of intercellular spaces. (The phyllids of the moss familyPolytrichaceae are notable exceptions.) The phyllids of bryophytes are only present on thegametophytes, while in contrast the leaves of vascular plants are only present on thesporophytes. These can further develop into either vegetative or reproductive structures.[13]
Simple, vascularized leaves (microphylls), such as those of the early Devonian lycopsidBaragwanathia, 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, inprogymnosperms likeArchaeopteris, inSphenopsida,ferns and later in thegymnosperms andangiosperms. Euphylls are also referred to as macrophylls or megaphylls (large leaves).[6]
A structurally complete leaf of anangiosperm consists of apetiole (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 astipe inferns. The lamina is the expanded, flat component of the leaf which contains thechloroplasts. The sheath is a structure, typically at the base that fully or partially clasps thestem above the node, where the leaf is attached. Leaf sheathes typically occur inPoaceae (grasses) andApiaceae (umbellifers). Between the sheath and the lamina, there may be apseudopetiole, a petiole like structure. Pseudopetioles occur in somemonocotyledons includingbananas,palms andbamboos.[17] Stipules may be conspicuous (e.g.beans androses), soon falling or otherwise not obvious as inMoraceae or absent altogether as in theMagnoliaceae. 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 abud. 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 orspecies levels, and botanists have developed a richterminology 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 orpolymorphic). The longest leaves are those of theRaffia palm,R. regalis which may be up to 25 m (82 ft) long and 3 m (9.8 ft) wide.[18] The terminology associated with the description of leaf morphology is presented, in illustrated form, atWikibooks.
Perennial plants whose leaves are shed annually are said to have deciduous leaves, while leaves that remain through winter areevergreens. Leaves attached to stems by stalks (known aspetioles) are called petiolate, and if attached directly to the stem with no petiole they are called sessile.[19]
Conifer leaves are typically needle- or awl-shaped or scale-like; they are usually evergreen but can sometimes be deciduous. Usually, they have a single vein.
The standard form of flowering plants (angiosperm) includesstipules, a petiole, and alamina.
Sheath leaves are the type found in mostgrasses and many other monocots.
Other specialized leaves include those ofNepenthes, a pitcher plant.
Dicot leaves have blades with pinnate venation (where major veins diverge from one large mid-vein and have smaller connecting networks between them). Less commonly, dicot leaf blades may have palmate venation (several large veins diverging frompetiole to leaf edges). Finally, some exhibit parallel venation.[19]Monocot leaves in temperate climates usually have narrow blades and usually parallel venation converging at leaf tips or edges. Some also have pinnate venation.[19]
The arrangement of leaves on the stem is known asphyllotaxis.[20] A large variety of phyllotactic patterns occur in nature:
The leaves on this plant are arranged in pairsopposite one another, with successive pairs at right angles to each other (decussate) along the red stem. Note the developing buds in the axils of these leaves.The leaves on this plant (Senecio angulatus) are alternately arranged.
Alternate
One leaf, branch, or flower part attaches at each point or node on the stem, and leaves alternate direction—to a greater or lesser degree—along the stem.
Basal
Arising from the base of the plant.
Cauline
Attached to the aerial stem.
Opposite
Two leaves, branches, or flower parts attach at each point or node on the stem. Leaf attachments are paired at each node.
Three or more leaves, branches, or flower parts 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.). Opposite leaves may appear whorled near the tip of the stem.Pseudoverticillate describes an arrangement only appearing whorled, but not actually so.
The termdistichous literally meanstwo rows. Leaves in this arrangement may be alternate or opposite in their attachment. The term2-ranked is equivalent. The termstristichous andtetrastichous are sometimes encountered. For example, the "leaves" (actually microphylls) of most species ofSelaginella are tetrastichous but not decussate.
In the simplest mathematical models of phyllotaxis, the apex of the stem is represented as a circle. Each new node is formed at the apex, and it is rotated by a constant angle from the previous node. This angle is called thedivergence angle. The number of leaves that grow from a node depends on the plant species. When a single leaf grows from each node, and when the stem is held straight, the leaves form ahelix.
The divergence angle is often represented as a fraction of a full rotation around the stem. A rotation fraction of 1/2 (a divergence angle of 180°) produces an alternate arrangement, such as inGasteria or the fan-aloeKumara plicatilis. Rotation fractions of 1/3 (divergence angles of 120°) occur inbeech andhazel.Oak andapricot rotate by 2/5, sunflowers, poplar, and pear by 3/8, and in willow and almond the fraction is 5/13.[21] These arrangements are periodic. Thedenominator of the rotation fraction indicates the number of leaves in one period, while thenumerator indicates the number of complete turns orgyres made in one period. For example:
180° (or1⁄2): two leaves in one circle (alternate leaves)
120° (or1⁄3): three leaves in one circle
144° (or2⁄5): five leaves in two gyres
135° (or3⁄8): eight leaves in three gyres.
Most divergence angles are related to the sequence ofFibonacci numbersFn. This sequence begins 1, 1, 2, 3, 5, 8, 13; each term is the sum of the previous two. Rotation fractions are often quotientsFn /Fn + 2 of a Fibonacci number by the number two terms later in the sequence. This is the case for the fractions 1/2, 1/3, 2/5, 3/8, and 5/13. The ratio between successive Fibonacci numbers tends to thegolden ratioφ = (1 + √5)/2. When a circle is divided into two arcs whose lengths are in the ratio1:φ, the angle formed by the smaller arc is thegolden angle, which is1/φ2 × 360° ≈ 137.5°. Because of this, many divergence angles are approximately137.5°. In plants where a pair of opposite leaves grows from each node, the leaves form a double helix. If the nodes do not rotate (a rotation fraction of zero and a divergence angle of 0°), the two helices become a pair of parallel lines, creating a distichous arrangement as inmaple orolive trees. More common in a decussate pattern, in which each node rotates by 1/4 (90°) as in the herbbasil. The leaves of tricussate plants such asNerium oleander form a triple helix. The leaves of some plants do not form helices. In some plants, the divergence angle changes as the plant grows.[22] In orixate phyllotaxis, named afterOrixa japonica, the divergence angle is not constant. Instead, it is periodic and follows the sequence 180°, 90°, 180°, 270°.[23]
Two basic forms of leaves can be described considering the way the blade (lamina) is divided. Asimple leaf has an undivided blade. However, the leaf may bedissected to form lobes, but the gaps between lobes do not reach to the main vein. Acompound leaf has a fully subdivided blade, eachleaflet of the blade being separated along a main or secondary vein. The leaflets may have petiolules and stipels, the equivalents of the petioles and stipules of leaves. 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 theFabaceae. The middle vein of a compound leaf or afrond, when it is present, is called arachis.
Palmately compound
The leaflets all have a common point of attachment at the end of the petiole, radiating like fingers of a hand; for example,Cannabis (hemp) andAesculus (buckeyes).
Pinnately compound
Leaflets are arranged either side of the main axis, orrachis.
Odd pinnate
With a terminal leaflet; for example,Fraxinus (ash).
Even pinnate
Lacking a terminal leaflet; for example,Swietenia (mahogany). A specific type of even pinnate isbifoliolate, where leaves only consist of two leaflets; for example,Hymenaea.
Bipinnately compound
Leaves are twice divided: the leaflets (technically "subleaflets") are arranged along a secondary axis that is one of several branching off the rachis. Each leaflet is called apinnule. The group of pinnules on each secondary vein forms apinna; for example,Albizia (silk tree).
Pinnately dissected to the central vein, but with the leaflets not entirely separate; for example,Polypodium, someSorbus (whitebeams). In pinnately veined leaves the central vein is known as themidrib.
The overgrown petioles ofrhubarb (Rheum rhabarbarum) are edible.
Leaves which have a petiole (leaf stalk) are said to bepetiolate.Sessile (epetiolate) leaves have no petiole, and the blade attaches directly to the stem. Subpetiolate leaves are nearly petiolate or have an extremely short petiole and may appear to be sessile. Inclasping ordecurrent leaves, the blade partially surrounds the stem. When the leaf base completely surrounds the stem, the leaves are said to beperfoliate, such as inEupatorium perfoliatum. In peltate leaves, the petiole attaches to the blade inside the blade margin. In someAcacia species, such as the koa tree (Acacia koa), the petioles are expanded or broadened and function like leaf blades; these are calledphyllodes. There may or may not be normal pinnate leaves at the tip of the phyllode. Astipule, present on the leaves of manydicotyledons, 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 inroses andbeans), 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".
Branching veins on underside oftaro leafThe venation within the bract of alindenMicrograph of a leaf skeleton
Veins (sometimes referred to as nerves) constitute one of the most visible features of leaves. The veins in a leaf represent the vascular structure of the organ, extending into the leaf via the petiole and providing transportation of water and nutrients between leaf and stem, and play a crucial role in the maintenance of leaf water status and photosynthetic capacity. They also play a role in the mechanical support of the leaf.[24][25] Within the lamina of the leaf, while some vascular plants possess only a single vein, in most this vasculature generally divides (ramifies) according to a variety of patterns (venation) and form cylindrical bundles, usually lying in the median plane of themesophyll, between the two layers ofepidermis.[26] This pattern is often specific to taxa, and of which angiosperms possess two main types,parallel andreticulate (net like). In general, parallel venation is typical of monocots, while reticulate is more typical ofeudicots andmagnoliids ("dicots"), though there are many exceptions.[27][26][28]
The vein or veins entering the leaf from the petiole are called primary or first-order veins. The veins branching from these are secondary or second-order veins. These primary and secondary veins are considered major veins or lower order veins, though some authors include third order.[29] Each subsequent branching is sequentially numbered, and these are the higher order veins, each branching being associated with a narrower vein diameter.[30]
In parallel veined leaves, the primary veins run parallel and equidistant to each other for most of the length of the leaf and then converge or fuse (anastomose) towards the apex. Usually, many smaller minor veins interconnect these primary veins but may terminate with very fine vein endings in the mesophyll. Minor veins are more typical of angiosperms, which may have as many as four higher orders.[29]
In contrast, leaves with reticulate venation have a single (sometimes more) primary vein in the centre of the leaf, referred to as the midrib or costa, which is continuous with the vasculature of the petiole. The secondary veins, also known as second order veins or lateral veins, branch off from the midrib and extend toward the leaf margins. These often terminate in ahydathode, a secretory organ, at the margin. In turn, smaller veins branch from the secondary veins, known as tertiary or third order (or higher order) veins, forming a dense reticulate pattern. The areas or islands of mesophyll lying between the higher order veins, are calledareoles. Some of the smallest veins (veinlets) may have their endings in the areoles, a process known as areolation.[30] These minor veins act as the sites of exchange between the mesophyll and the plant's vascular system.[25] Thus, minor veins collect the products of photosynthesis (photosynthate) from the cells where it takes place, while major veins are responsible for its transport outside of the leaf. At the same time water is being transported in the opposite direction.[31][27][26]
The number of vein endings is variable, as is whether second order veins end at the margin, or link back to other veins.[28] There are many elaborate variations on the patterns that the leaf veins form, and these have functional implications. Of these, angiosperms have the greatest diversity.[29] Within these the major veins function as the support and distribution network for leaves and are correlated with leaf shape. For instance, the parallel venation found in most monocots correlates with their elongated leaf shape and wide leaf base, while reticulate venation is seen in simple entire leaves, while digitate leaves typically have venation in which three or more primary veins diverge radially from a single point.[32][25][30][33]
In evolutionary terms, early emerging taxa tend to have dichotomous branching with reticulate systems emerging later. Veins appeared in thePermian, prior to the appearance of angiosperms in theTriassic, during which vein hierarchy appeared enabling higher function, larger leaf size and adaption to a wider variety of climatic conditions.[29] Although it is the more complex pattern, branching veins appear to beplesiomorphic and in some form were present in ancientseed plants as long as 250 million years ago. A pseudo-reticulate venation that is actually a highly modified penniparallel one is anautapomorphy of someMelanthiaceae, which are monocots; e.g.,Paris quadrifolia (True-lover's Knot). In leaves with reticulate venation, veins form a scaffolding matrix imparting mechanical rigidity to leaves.[34]
Leaves are normally extensively vascularized and typically have networks ofvascular bundles containingxylem, which supplies water forphotosynthesis, andphloem, which transports thesugars produced by photosynthesis. Many leaves are covered intrichomes (small hairs) which have diverse structures and functions.
These three tissue systems typically form a regular organization at the cellular scale. Specialized cells that differ markedly from surrounding cells, and which often synthesize specialized products such as crystals, are termedidioblasts.[35]
SEM image of the leaf epidermis ofNicotiana alata, showingtrichomes (hair-like appendages) andstomata (eye-shaped slits, visible at full resolution).
Theepidermis is the outer layer ofcells covering the leaf. It is covered with a waxycuticle which is impermeable to liquid water and water vapor and forms the boundary separating the plant's inner cells from the external world. The cuticle is in some cases thinner on the lower epidermis than on the upper epidermis, and is generally thicker on leaves from dry climates as compared with those from wet climates.[36] The epidermis serves several functions: protection against water loss by way oftranspiration, regulation of gas exchange and secretion ofmetabolic compounds. Most leaves show dorsoventral anatomy: The upper (adaxial) and lower (abaxial) surfaces have somewhat different construction and may serve different functions.
The epidermis tissue includes several differentiated cell types; epidermal cells, epidermal hair cells (trichomes), cells in the stomatal complex; guard cells and subsidiary cells. The epidermal cells are the most numerous, largest, and least specialized and form the majority of the epidermis. They are typically more elongated in the leaves ofmonocots than in those ofdicots.
Chloroplasts are generally absent in epidermal cells, the exception being the guard cells of thestomata. The stomatal pores perforate the epidermis and are surrounded on each side by chloroplast-containing guard cells, and two to four subsidiary cells that lack chloroplasts, forming a specialized cell group known as the stomatal complex. The opening and closing of the stomatal aperture is controlled by the stomatal complex and regulates the exchange of gases and water vapor between the outside air and the interior of the leaf. Stomata therefore play the important role in allowing photosynthesis without letting the leaf dry out. In a typical leaf, the stomata are more numerous over the abaxial (lower) epidermis than the adaxial (upper) epidermis and are more numerous in plants from cooler climates.
For the termMesophyll in the size classification of leaves, seeLeaf size.
Most of the interior of the leaf between the upper and lower layers of epidermis is aparenchyma (ground tissue) orchlorenchyma tissue called themesophyll (Greek for "middle leaf"). Thisassimilation 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:
An upperpalisade layer of vertically elongated cells, one to two cells thick, directly beneath the adaxial epidermis, with intercellular air spaces between them. Its cells contain many more chloroplasts than the spongy layer. Cylindrical cells, with thechloroplasts close to the walls of the cell, can take optimal advantage of light. The slight separation of the cells provides maximumabsorption of carbon dioxide. 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 thespongy layer. The cells of the spongy layer are more branched and not so tightly packed, so that there are large intercellular air spaces between them. The pores orstomata of the epidermis open into substomatal chambers, which are connected to the intercellular air spaces between the spongy and palisade mesophyll cell, so that oxygen, carbon dioxide and water vapor can diffuse into and out of the leaf and access the mesophyll cells during respiration, photosynthesis and transpiration.
Leaves are normally green, due to chlorophyll in chloroplasts in the mesophyll cells. Some plants have leaves of different colours due to the presence ofaccessory pigments such ascarotenoids in their mesophyll cells.
Cells that usually movesap, with dissolved sucrose (glucose to sucrose) produced by photosynthesis in the leaf, out of the leaf.
The xylem typically lies on the adaxial side of the vascular bundle and the phloem typically lies on the abaxial side. Both are embedded in a dense parenchyma tissue, called the sheath, which usually includes some structural collenchyma tissue.
According toAgnes Arber's partial-shoot theory of the leaf, leaves are partial shoots,[40] being derived from leafprimordia of the shoot apex. Early in development they are dorsiventrally flattened with both dorsal and ventral surfaces.[13] Compound leaves are closer to shoots than simple leaves. Developmental studies have shown that compound leaves, like shoots, may branch in three dimensions.[41][42] On the basis of molecular genetics, Eckardt and Baum (2010) concluded that "it is now generally accepted that compound leaves express both leaf and shoot properties."[43] Many dicotyledonous leaves show endogenously driven daily rhythmicity in growth.[44][45][46]
Plants respond and adapt to environmental factors, such as light and mechanical stress from wind. Leaves need to support their own mass and align themselves in such a way as to optimize their exposure to the sun, generally more or less horizontally. However, horizontal alignment maximizes exposure to bending forces and failure from stresses such as wind, snow, hail, falling debris, animals, and abrasion from surrounding foliage and plant structures. Overall leaves are relatively flimsy with regard to other plant structures such as stems, branches and roots.[47]
Both leaf blade and petiole structure influence the leaf's response to forces such as wind, allowing a degree of repositioning to minimizedrag and damage, as opposed to resistance. Leaf movement like this may also increaseturbulence of the air close to the surface of the leaf, which thins theboundary layer of air immediately adjacent to the surface, increasing the capacity for gas and heat exchange, as well as photosynthesis. Strong wind forces may result in diminished leaf number and surface area, which while reducing drag, involves atrade off of also reducing photosynthesis. Thus, leaf design may involve compromise between carbon gain, thermoregulation and water loss on the one hand, and the cost of sustaining both static and dynamic loads. In vascular plants, perpendicular forces are spread over a larger area and are relatively flexible in both bending andtorsion, enabling elastic deforming without damage.[47]
Many leaves rely onhydrostatic support arranged around a skeleton of vascular tissue for their strength, which depends on maintaining leaf water status. Both the mechanics and architecture of the leaf reflect the need for transportation and support. Read and Stokes (2006) consider two basic models, the "hydrostatic" and "I-beam leaf" form (see Fig 1).[47] Hydrostatic leaves such as inProstanthera lasianthos are large and thin, and may involve the need for multiple leaves rather single large leaves because of the amount of veins needed to support the periphery of large leaves. But large leaf size favors efficiency in photosynthesis and water conservation, involving further trade offs. On the other hand, I-beam leaves such asBanksia marginata involve specialized structures to stiffen them. These I-beams are formed from bundle sheath extensions ofsclerenchyma meeting stiffened sub-epidermal layers. This shifts the balance from reliance on hydrostatic pressure to structural support, an obvious advantage where water is relatively scarce.[47] Long narrow leaves bend more easily than ovate leaf blades of the same area. Monocots typically have such linear leaves that maximize surface area while minimising self-shading. In these a high proportion of longitudinal main veins provide additional support.[47]
Although not as nutritious as other organs such as fruit, leaves provide a food source for many organisms. The leaf is a vital source of energy production for the plant, and plants have evolved protection against animals that consume leaves, such astannins, chemicals which hinder the digestion of proteins and have an unpleasant taste. Animals that are specialized to eat leaves are known asfolivores.
Some species havecryptic adaptations by which they use leaves in avoiding predators. For example, the caterpillars ofsome leaf-roller moths will create a small home in the leaf by folding it over themselves. Several otherlepidopteran larvae modify leaves for shelter; perhaps the greatest variety of shelter types occurs among theskipper butterflies (Hesperiidae), which will cut, fold, and bind leaves usingsilk.[48] Somesawflies similarly roll the leaves of their food plants into tubes. Females of theAttelabidae, so-called leaf-rolling weevils, lay their eggs into leaves that they then roll up as means of protection. Other herbivores and their predatorsmimic the appearance of the leaf. Reptiles such as some chameleons, and insects such as somekatydids, also mimic the oscillating movements of leaves in the wind, moving from side to side or back and forth while evading a possible threat.
Leaves intemperate,boreal, and seasonally dry zones may be seasonally deciduous (falling off or dying for the inclement season). This mechanism to shed leaves is calledabscission. When the leaf is shed, it leaves a leaf scar on the twig. In cold autumns, they sometimeschange color, and turnyellow, bright-orange, orred, as various accessory pigments (carotenoids andxanthophylls) are revealed when the tree responds to cold and reducedsunlight by curtailing chlorophyll production. Redanthocyanin pigments are now thought to be produced in the leaf as it dies, possibly to mask the yellow hue left when the chlorophyll is lost—yellow leaves appear to attract herbivores such asaphids.[49] Optical masking of chlorophyll by anthocyanins reduces risk of photo-oxidative damage to leaf cells as they senesce, which otherwise may lower the efficiency of nutrient retrieval from senescing autumn leaves.[50]
Poinsettiabracts are leaves which have evolved red pigmentation in order to attract insects and birds to the central flowers, an adaptive function normally served bypetals (which are themselves leaves highly modified by evolution).
Large surface area provides a large area for capture of sunlight.
In harmful levels of sunlight, specialized leaves, opaque or partly buried, admit light through a translucentleaf window for photosynthesis at inner leaf surfaces (e.g.Fenestraria).
Special leaves on carnivorous plants are adapted for trapping food, mainly invertebrate prey, though some species trap small vertebrates as well (seecarnivorous plants).
Common mullein (Verbascum thapsus) leaves are covered in dense, stellate trichomes.Scanning electron microscope image of trichomes on the lower surface of aColeus blumei (coleus) leafSilky aster (Symphyotrichum sericeum) leaves are sericeous.
"Hairs" on plants are properly calledtrichomes. Leaves can show several degrees of hairiness. The meaning of several of the following terms can overlap.
Arachnoid, or arachnose
With many fine, entangled hairs giving a cobwebby appearance.
A number of different classification systems of the patterns of leaf veins (venation or veination) have been described,[28] starting with Ettingshausen (1861),[53] together with many different descriptive terms, and the terminology has been described as "formidable".[28] One of the commonest among these is the Hickey system, originally developed for "dicotyledons" and using a number of Ettingshausen's terms derived from Greek (1973–1979):[54][55][56] (see also: Simpson Figure 9.12, p. 468)[28]
1.Pinnate (feather-veined, reticulate, pinnate-netted, penniribbed, penninerved, or penniveined)
The veins arisepinnately (feather like) from a single primary vein (mid-vein) and subdivide into secondary veinlets, known as higher order veins. These, in turn, form a complicated network. This type of venation is typical for (but by no means limited to) "dicotyledons" (non monocotyledonangiosperms). E.g.,Ostrya. There are three subtypes of pinnate venation:
Major veins extend close to the margin, but bend before they intersect with the margin.
Hyphodromous
All secondary veins are absent, rudimentary or concealed
These in turn have a number of further subtypes such as eucamptodromous, where secondary veins curve near the margin without joining adjacent secondary veins.
Two or more primary veins originating beside each other at the leaf base, and runningparallel to each other to the apex and then converging there. Commissural veins (small veins) connect the major parallel veins. Typical for mostmonocotyledons, such asgrasses. The additional terms marginal (primary veins reach the margin), and reticulate (net-veined) are also used.
Parallelodromous
3. Campylodromous (campylos – curve)
Several primary veins or branches originating at or close to a single point and running in recurved arches, then converging at apex. E.g.Maianthemum .
Campylodromous
4. Acrodromous
Two or more primary or well developed secondary veins in convergent arches towards apex, without basal recurvature as in Campylodromous. May be basal or suprabasal depending on origin, and perfect or imperfect depending on whether they reach to 2/3 of the way to the apex. E.g.,Miconia (basal type),Endlicheria (suprabasal type).
Acrodromous
Imperfect basal
Imperfect suprabasal
Perfect basal
Perfect suprabasal
5. Actinodromous
Three or more primary veins diverging radially from a single point. E.g.,Arcangelisia (basal type),Givotia (suprabasal type).
Primary veins with one or more points of secondary dichotomous branching beyond the primary divergence, either closely or more distantly spaced. E.g.,Platanus.
Venation of a Poinsettia (Euphorbia pulcherrima) leaf.
Palinactodromous
Types 4–6 may similarly be subclassified as basal (primaries joined at the base of the blade) or suprabasal (diverging above the blade base), and perfect or imperfect, but also flabellate.
At about the same time, Melville (1976) described a system applicable to all Angiosperms and using Latin and English terminology.[57] Melville also had six divisions, based on the order in which veins develop.
Arbuscular (arbuscularis)
Branching repeatedly by regular dichotomy to give rise to a three dimensional bush-like structure consisting of linear segment (2 subclasses)
Flabellate (flabellatus)
Primary veins straight or only slightly curved, diverging from the base in a fan-like manner (4 subclasses)
Palmate (palmatus)
Curved primary veins (3 subclasses)
Pinnate (pinnatus)
Single primary vein, the midrib, along which straight or arching secondary veins are arranged at more or less regular intervals (6 subclasses)
Collimate (collimatus)
Numerous longitudinally parallel primary veins arising from a transverse meristem (5 subclasses)
Conglutinate (conglutinatus)
Derived from fused pinnate leaflets (3 subclasses)
A modified form of the Hickey system was later incorporated into the Smithsonian classification (1999) which proposed seven main types of venation, based on the architecture of the primary veins, adding Flabellate as an additional main type. Further classification was then made on the basis of secondary veins, with 12 further types, such as;
Brochidodromous
Closed form in which the secondaries are joined in a series of prominent arches, as inHildegardia.
Craspedodromous
Open form with secondaries terminating at the margin, in toothed leaves, as inCeltis.
Eucamptodromous
Intermediate form with upturned secondaries that gradually diminish apically but inside the margin, and connected by intermediate tertiary veins rather than loops between secondaries, as inCornus.
Cladodromous
Secondaries freely branching toward the margin, as inRhus.
terms which had been used as subtypes in the original Hickey system.[58]
Several to many equal fine basal veins diverging radially at low angles and branching apically. E.g.Paranomus.
Flabellate
Analyses of vein patterns often fall into consideration of the vein orders, primary vein type, secondary vein type (major veins), and minor vein density. A number of authors have adopted simplified versions of these schemes.[59][28] At its simplest the primary vein types can be considered in three or four groups depending on the plant divisions being considered;
pinnate
palmate
parallel
where palmate refers to multiple primary veins that radiate from the petiole, as opposed to branching from the central main vein in the pinnate form, and encompasses both of Hickey types 4 and 5, which are preserved as subtypes; e.g., palmate-acrodromous (see National Park Service Leaf Guide).[60]
Several main veins of approximately equal sizediverge from a common point near the leaf base where the petiole attaches, and radiate toward the edge of the leaf. Palmately veined leaves are often lobed or divided with lobes radiating from the common point. They may vary in the number of primary veins (3 or more), but always radiate from a common point.[61] e.g. mostAcer (maples).
Veins successively branching into equally sized veins from a common point, forming a Y junction, fanning out. Amongst temperate woody plants,Ginkgo biloba is the only species exhibiting dichotomous venation. Also somepteridophytes (ferns).[61]
Parallel
Primary and secondary veins roughly parallel to each other, running the length of the leaf, often connected by short perpendicular links, rather than form networks. In some species, the parallel veins join at the base and apex, such as needle-type evergreens and grasses. Characteristic of monocotyledons, but exceptions includeArisaema, and as below, under netted.[61]
Netted (reticulate, pinnate)
A prominent midvein with secondary veins branching off along both sides of it. The name derives from the ultimate veinlets which form an interconnecting net like pattern or network. (The primary and secondary venation may be referred to as pinnate, while the net like finer veins are referred to as netted or reticulate); most non-monocot angiosperms, exceptions includingCalophyllum. Some monocots have reticulate venation, includingColocasia,Dioscorea andSmilax.[61]
Equisetum: Reduced microphyllous leaves (L) arising in whorl from node
However, these simplified systems allow for further division into multiple subtypes. Simpson,[28] (and others)[62] divides parallel and netted (and some use only these two terms for Angiosperms)[63] on the basis of the number of primary veins (costa) as follows;
Single central prominent midrib, secondary veins from this arise perpendicularly to it and run parallel to each other towards the margin or tip, but do not join (anastomose). The term unicostate refers to the prominence of the single midrib (costa) running the length of the leaf from base to apex. e.g.Zingiberales, such asBananas etc.
Palmate-parallel (multicostate parallel)
Several equally prominent primary veins arising from a single point at the base and running parallel towards tip or margin. The term multicostate refers to having more than one prominent main vein. e.g."fan" (palmate) palms (Arecaceae)
Mid-veins diverge more or less parallel towards the margin e.g.Borassus (Poaceae), fan palms
Netted (Reticulate)
Pinnately (veined, netted, unicostate reticulate)
Single prominent midrib running from base to apex, secondary veins arising on both sides along the length of the primary midrib, running towards the margin or apex (tip), with a network of smaller veinlets forming a reticulum (mesh or network). e.g.Mangifera,Ficus religiosa,Psidium guajava,Hibiscus rosa-sinensis,Salix alba
Palmately (multicostate reticulate)
More than one primary veins arising from a single point, running from base to apex. e.g.Liquidambar styraciflua This may be further subdivided;
Multicostate convergent
Major veins diverge from origin at base then converge towards the tip. e.g.Zizyphus,Smilax,Cinnamomum
These complex systems are not used much in morphological descriptions of taxa, but have usefulness in plant identification,[28] although criticized as being unduly burdened with jargon.[66]
An older, even simpler system, used in some flora[67] uses only two categories, open and closed.
Open: Higher order veins have free endings among the cells and are more characteristic of non-monocotyledon angiosperms. They are more likely to be associated with leaf shapes that are toothed, lobed or compound. They may be subdivided as;
Pinnate (feather-veined) leaves, with a main central vein or rib (midrib), from which the remainder of the vein system arises
Palmate, in which three or more main ribs rise together at the base of the leaf, and diverge upward.
Dichotomous, as in ferns, where the veins fork repeatedly
Closed: Higher order veins are connected in loops without ending freely among the cells. These tend to be in leaves with smooth outlines, and are characteristic of monocotyledons.
They may be subdivided into whether the veins run parallel, as in grasses, or have other patterns.
There are also many other descriptive terms, often with very specialized usage and confined to specific taxonomic groups.[68] The conspicuousness of veins depends on a number of features. These include the width of the veins, their prominence in relation to the lamina surface and the degree of opacity of the surface, which may hide finer veins. In this regard, veins are calledobscure and the order of veins that are obscured and whether upper, lower or both surfaces, further specified.[69][61]
Terms that describe vein prominence includebullate,channelled,flat,guttered,impressed,prominent andrecessed (Fig. 6.1 Hawthorne & Lawrence 2013).[66][70] Veins may show different types of prominence in different areas of the leaf. For instancePimenta racemosa has a channelled midrib on the upper surface, but this is prominent on the lower surface.[66]
Describing vein prominence:
Bullate
Surface of leaf raised in a series of domes between the veins on the upper surface, and therefore also with marked depressions. e.g.Rytigynia pauciflora,[71]Vitis vinifera
Channelled (canalicululate)
Veins sunken below the surface, resulting in a rounded channel. Sometimes confused with "guttered" because the channels may function as gutters for rain to run off and allow drying, as in manyMelastomataceae.[72] e.g. (see)Pimenta racemosa (Myrtaceae),[73]Clidemia hirta (Melastomataceae).
Guttered
Veins partly prominent, the crest above the leaf lamina surface, but with channels running along each side, like gutters
Impressed
Vein forming raised line or ridge which lies below the plane of the surface which bears it, as if pressed into it, and are often exposed on the lower surface. Tissue near the veins often appears to pucker, giving them a sunken or embossed appearance
Obscure
Veins not visible, or not at all clear; if unspecified, then not visible with the naked eye. e.g.Berberis gagnepainii. In thisBerberis, the veins are only obscure on the undersurface.[74]
More than one main vein (nerve) at the base. Lateral secondary veins branching from a point above the base of the leaf. Usually expressed as asuffix, as in 3-plinerved or triplinerved leaf. In a 3-plinerved (triplinerved) leaf three main veins branch above the base of the lamina (two secondary veins and the main vein) and run essentially parallel subsequently, as inCeanothus and inCeltis. Similarly, a quintuplinerve (five-veined) leaf has four secondary veins and a main vein. A pattern with 3–7 veins is especially conspicuous inMelastomataceae. The term has also been used inVaccinieae. The term has been used as synonymous with acrodromous, palmate-acrodromous or suprabasal acrodromous, and is thought to be too broadly defined.[76][76]
Scalariform
Veins arranged like the rungs of a ladder, particularly higher order veins
The termsmegaphyll,macrophyll,mesophyll,notophyll,microphyll,nanophyll andleptophyll are used to describe leaf sizes (in descending order), in a classification devised in 1934 byChristen C. Raunkiær and since modified by others.[77][78]
^abcBotany Illustrated: Introduction to Plants Major Groups Flowering Plant Families. Thomson Science. 1984. p. 21.
^Didier Reinhardt and Cris Kuhlemeier, "Phyllotaxis in higher plants", in Michael T. McManus, Bruce Veit, eds.,Meristematic Tissues in Plant Growth and Development, January 2002,ISBN978-1-84127-227-6, Wiley-Blackwell.
^Coxeter HS (1961).Introduction to geometry. Wiley. p. 169.
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