A leaf (pl: leaves) is any of the principal appendages of a vascular plant stem,[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 that 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 includes 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.

Contents

1 General characteristics

2 Morphology

2.1 Basic leaf types

2.2 Arrangement on the stem

2.3 Divisions of the blade

2.4 Characteristics of the petiole

2.5 Veins

2.6 Morphology changes within a single plant

3 Anatomy
3.1 Medium-scale features

3.2 Small-scale features

3.3 Major leaf tissues

3.3.1 Epidermis

3.3.2 Mesophyll

3.3.3 Vascular tissue

4 Leaf development

5 Ecology

5.1 Biomechanics

5.2 Interactions with other organisms

5.3 Seasonal leaf loss

6 Evolutionary adaptation

7 Terminology

7.1 Shape

7.2 Edge (margin)

7.3 Apex (tip)

7.4 Base

7.5 Surface

7.6 Hairiness

7.7 Timing

7.8 Venation

7.8.1 Classification

7.8.1.1 Hickey system

7.8.1.2 Other systems

7.8.2 Other descriptive terms

7.8.3 Diagrams of venation patterns

7.9 Size

8 See also

9 References
10 Bibliography

10.1 Books and chapters

10.2 Articles and theses

10.3 Websites

11 External links

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 

Vein skeleton of a leaf. Veins contain lignin that make them harder to degrade for microorganisms.

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, oxygen, 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]

Near the ground these Eucalyptus 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 (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).