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From Wikipedia, the free encyclopedia

The leaves of a Beech tree

In botany, a leaf is an above-ground plant organ specialized for photosynthesis. For this purpose, a leaf is typically flat (laminar) and thin. There is continued debate about whether the flatness of leaves evolved to expose the chloroplasts to more light or to increase the absorption of carbon dioxide. In either case, the adaption was made at the expense of water loss. In the Devonian period, when carbon dioxide concentration was at several times its present value, plants did not have leaves or flat stems. Many bryophytes have flat, photosynthetic organs, but these are not true leaves. Neither are the microphylls of lycophytes. The leaves of ferns, gymnosperms, and angiosperms are variously referred to as macrophyll, megaphylls, or euphylls.

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.

Contents

Anatomy

Cross section of a leaf

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 aforementioned structural components. In certain 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. Periodically (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:

  1. An epidermis that covers the upper and lower surfaces
  2. An interior chlorenchyma called the mesophyll
  3. An arrangement of veins (the vascular tissue)
Diagram of leaf internal anatomy

Epidermis

Epidermal cells

The epidermis is the outer layer 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.

SEM image of Nicotiana alata leaf's epidermis, showing trichomes (hair-like appendages) and stomata (eye-shaped slits, visible at full resolution).

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.

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 (Greek for "middle leaf"). This assimilation tissue is the primary location of photosynthesis in the plant. The products of photosynthesis are called "assimilates".

Palisade cells

In ferns and most flowering plants the mesophyll 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 maximum 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.
Spongy cells
  • 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. These cells contain fewer chloroplasts than those of the palisade layer.

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 are normally green in color, which comes from chlorophyll found in plastids in the chlorenchyma cells. Plants that lack chlorophyll cannot photosynthesize.

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, possibly to mask the yellow hue left when the chlorophyll is lost - yellow leaves appear to attract herbivores such as aphids.[1]

Veins

Vein skeleton of a Hydrangea leaf

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:

  • Xylem: tubes that brings water and minerals from the roots into the leaf.
  • Phloem: tubes that usually move sap, with dissolved sucrose, produced by photosynthesis in the leaf, out of the leaf.

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

Morphology

The Citrus leaf is identified by the pores and pigments, as well as the margins.

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

Leaves of the White Spruce (Picea glauca) are needle-shaped and their arrangement is spiral

Arrangement on the stem

Different terms are usually used to describe leaf placement (phyllotaxis):

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.
  • Alternate — leaf attachments are singular at nodes, and leaves alternate direction, to a greater or lesser degree, along the stem.
  • Opposite — leaf attachments are paired at each node; decussate if, as typical, each successive pair is rotated 90° progressing along the stem; or distichous if not rotated, but two-ranked (in the same geometric flat-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). Opposite leaves may appear whorled near the tip of the stem.
  • Rosulate — leaves form a rosette

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 close to 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:

  • alternate leaves have an angle of 180° (or 1/2)
  • 120° (or 1/3) : three leaves in one circle
  • 144° (or 2/5) : five leaves in two gyres
  • 135° (or 3/8) : eight leaves in three gyres.

Divisions of the lamina (blade)

A leaf with laminar structure and pinnate venation

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.

  • Palmately compound leaves have the leaflets radiating from the end of the petiole, like fingers off the palm of a hand, e.g. Cannabis (hemp) and Aesculus (buckeyes).
  • Pinnately compound leaves have the leaflets arranged along the main or mid-vein.
    • 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 (or trifoliolate): a pinnate leaf with just three leaflets, e.g. Trifolium (clover), Laburnum (laburnum).
  • pinnatifid: pinnately dissected to the central vein, but with the leaflets not entirely separate, e.g. Polypodium, some Sorbus (whitebeams). In pinnately veined leaves the central vein in known as the midrib.

Characteristics of the petiole

The overgrown petioles of Rhubarb (Rheum rhabarbarum) are edible.

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).

  • 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

Venation (arrangement of the veins)

Branching veins on underside of taro leaf
The venation within the bract of a Lime tree.
The lower epidermis of Tilia x europea
Palmate-veined leaf

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.

  • 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 (but by no means limited to) 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 branch at the base of the lamina and run essentially parallel subsequently, as in Ceanothus. A similar pattern (with 3-7 veins) is especially conspicuous in Melastomataceae.
    • 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 for 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.

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).

Morphology changes within a single plant

  • Homoblasty - Characteristic in which a plant has small changes in leaf size, shape, and growth habit between juvenile and adult stages.
  • Heteroblasty - Characteristic in which a plant has marked changes in leaf size, shape, and growth habit between juvenile and adult stages.


Terminology

Chart illustrating some leaf morphology terms
A portion of a celery leaf

Shape

Margins (edge)

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

  • ciliate: fringed with hairs
  • crenate: wavy-toothed; dentate with rounded teeth, such as Fagus (beech)
  • crenulate finely or shallowly crenate
  • 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)
  • entire: even; with a smooth margin; without toothing
  • 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).

Tip of the leaf

Leaves showing various morphologies. Clockwise from upper left: tripartite lobation, elliptic with serrulate margin, peltate with palmate venation, acuminate odd-pinnate (center), pinnatisect, lobed, elliptic with entire margin
  • 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.

Base of the leaf

  • 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 notch towards the stalk.
  • 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 shape.
  • sagittate: shaped like an arrowhead and with the acute basal lobes pointing downward.
  • truncate: ending abruptly with a flat end, that looks cut off.

Surface of the leaf

Scale-shaped leaves of a Norfolk Island Pine, Araucaria heterophylla.

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, or 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, or viscous: covered with thick, sticky secretions.

The leaf surface is also host to a large variety of microorganisms; in this context it is referred to as the phyllosphere.

The parallel veins within an iris leaf.

Hairiness (trichomes)

Common Mullein (Verbascum thapsus) leaves are covered in dense, stellate trichomes.
Scanning electron microscope image of trichomes on the lower surface of a Coleus blumei (coleus) leaf.

"Hairs" on plants are properly called trichomes. 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.
  • 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.
  • glabrous: no hairs of any kind present.
  • 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, or lanose: with woolly hairs.
  • pilose: with soft, clearly separated hairs.
  • puberulent, or puberulous: with fine, minute hairs.
  • pubescent: with soft, short and erect hairs.
  • scabrous, or 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, or stelliform: 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.

Adaptations

Poinsettia bracts are leaves which have evolved red pigmentation in order to attract insects and birds to the central flowers, an adaptive function normally served by petals (which are themselves leaves highly modified by evolution).

In the course of evolution, leaves have adapted to different environments in the following ways:

  • A certain surface structure avoids moistening by rain and contamination (See Lotus effect).
  • Sliced leaves reduce wind resistance.
  • Hairs on the leaf surface trap humidity in dry climates and create a large boundary layer thereby reducing water loss.
  • Waxy leaf surfaces reduce water loss.
  • Large surface area of leaf provides large area for sunlight and provides shade for plant to minimize heating and reduce water loss.
  • In more or less opaque or buried in the soil leaves, translucent windows filter the light before the photosynthesis takes place at the inner leaf surfaces (e.g. Fenestraria).
  • Succulent leaves store water and organic acids for use in CAM photosynthesis.
  • Aromatic oils, poisons or pheromones produced by leaf borne glands deter herbivores (e.g. eucalypts).
  • Inclusions of crystalline minerals deter herbivores (e.g. silica in grasses.
  • A transformation into petals attracts pollinators.
  • A transformation into spines protects the plants (e.g. cacti).
  • A transformation into insect traps helps feeding the plants (carnivorous plants).
  • A transformation into bulbs helps storing food and water (e.g. onions).
  • A transformation into tendrils allows the plant to climb (e.g. peas).
  • A transformation into bracts and pseudanthia (false flowers) replaces normal flower structures if the true flowers are extremely reduced (e.g. Spurges).

Interactions with other organisms

Leaf insects mimic leaves (Kallima inachus shown)

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.

Bibliography

  • Leaves: The formation, charactistics and uses of hundred of leaves in all parts of the world by Ghillean Tolmie Prance. 324 photographic plates in black and white, and colour by Kjell B Sandved 256 pages[2]

Footnotes

  1. ^ Thomas F. Döring; Marco Archetti; Jim Hardie (2009), "Autumn leaves seen through herbivore eyes" ( – Scholar search), Proceedings of the Royal Society B Biological Sciences 276: 121, doi:10.1098/rspb.2008.0858, http://users.ox.ac.uk/~zool0643/papers/PRSB_2008_silwood.pdf 
  2. ^ Published by Thames and Hudson (London) with an ISBN 0 500 54104 3

See also

External links



Source material

Up to date as of January 22, 2010
(Redirected to A Leaf article)

From Wikisource

A Leaf
by Ella Wheeler Wilcox
From Poems of Cheer (1910)

Somebody said, in the crowd, last eve,
   That you were married, or soon to be.
I have not thought of you, I believe,
   Since last we parted. Let me see:
Five long Summers have passed since then -
   Each has been pleasant in its own way -
And you are but one of a dozen men
   Who have played the suitor a Summer day.

But, nevertheless, when I heard your name,
   Coupled with some one's, not my own,
There burned in my bosom a sudden flame,
   That carried me back to the day that is flown.
I was sitting again by the laughing brook,
   With you at my feet, and the sky above,
And my heart was fluttering under your look -
   The unmistakable look of Love.

Again your breath, like a South wind, fanned
   My cheek, where the blushes came and went;
And the tender clasp of your strong, warm hand
   Sudden thrills through my pulses sent.
Again you were mine by Love's own right -
   Mine for ever by Love's decree:
So for a moment it seemed last night,
   When somebody mentioned your name to me.

Just for the moment I thought you mine -
   Loving me, wooing me, as of old.
The tale remembered seemed half divine -
   Though I held it lightly enough when told.
The past seemed fairer than when it was near,
   As "blessings brighten when taking flight;"
And just for the moment I held you dear -
   When somebody mentioned your name last night.

PD-icon.svg This work is in the public domain in the United States because it was published before January 1, 1923.

The author died in 1919, so this work is also in the public domain in countries and areas where the copyright term is the author's life plus 80 years or less. This work may also be in the public domain in countries and areas with longer native copyright terms that apply the rule of the shorter term to foreign works.


1911 encyclopedia

Up to date as of January 14, 2010

From LoveToKnow 1911

LEAF (0. Eng. leaf, cf. Dutch loof, Ger. Laub, Swed. lof, &c.; possibly to be referred to the root seen in Gr. X irECV, to peel, strip), the name given in popular language to all the green expanded organs borne upon an axis, and so applied to similar objects, such as a thin sheet of metal, a hinged flap of a table, the page of a book, &c.

Table of contents

Definition

Investigation has shown that many other parts of a plant which externally appear very different from ordinary leaves are, in their essential particulars, very similar to them, and are in fact their morphological equivalents. Such are the scales of a bulb, and the various parts of the flower, and assuming that the structure ordinarily termed a leaf is the typical form, these other structures were designated changed or metamorphosed leaves, a somewhat misleading interpretation. All structures morphologically equivalent with the leaf are now included under the general term phyllome (leaf-structure).

Leaves are produced as lateral outgrowths of the stem in definite succession below the apex. This character, common to all leaves, distinguishes them from other organs. In the higher plants we can easily recognize the distinction between stem and leaf. Amongst the lower plants, however, it is found that a demarcation into stem and leaf is impossible, but that there is a structure which partakes of the characters of both - such is a thallus. The leaves always arise from the outer portion of the primary meristem of the plant, and the tissues of the leaf are continuous with those of the stem. Every leaf originates as a simple cellular papilla (fig 1), which consists of a development from the cortical layers covered by epidermis; and as growth proceeds, the fibro-vascular bundles of the stem are continued outwards, and finally expand and terminate in the leaf. The increase in length of the leaf by growth at the apex is usually of a limited nature. In some ferns, however, there seems to be a provision for indefinite terminal growth, while in others this, growth is periodically interrupted. It not unfrequently happens, especially amongst Monocotyledons, that after growth at the apex has ceased, it is continued at the base of the leaf, and in this way the length may be much increased. Amongst Dicotyledons this is very rare. In all cases the dimensions of the leaf are enlarged by interstitial growth of its parts.

Parts of Leaves

The simplest leaf is found in some mosses, where it consists of a single layer of cells. The typical foliage leaf consists of several layers, and amongst vascular plants is distinguishable into an outer layer (epidermis) and a central tissue (parenchyma) with fibro-vascular bundles distributed through it.

Epidermis

The epidermis (fig. 2, es, ei), composed of cells more or less compressed, has usually a different structure and aspect on the two surfaces of the leaf. The cells of the epidermis are very closely united laterally and contain no green colouring matter (chlorophyll) except in the pair of cells - guard-cells - which bound the stomata. The outer wall, especially of the upper epidermis, has a tough outer layer or cuticle which renders it impervious to water. The epidermis is continuous except where stomata or spaces bounded by specialized cells communicate with intercellular spaces in the interior of the leaf. It is chiefly on the epidermis of the lower surface (fig. 2, ei) that stomata, st, are produced, and it is there also that hairs, p, usually occur. The lower epidermis is often of a dull or and pale-green colour. The upper epidermis is frequently smooth and shining, and sometimes becomes very hard and dense. Many tropical plants present on the upper surface of their leaves several layers of compressed cells beneath the epidermis which serve for storage of water and are known as aqueous tissue. In leaves which float upon the surface of the water, as those of the water-lily stomata.

Parenchyma and Mesophyll

The parenchyma of the leaf is the cellular tissue enclosed within the epidermis and surrounding the vessels (fig. 2, ps, pi). It is known as mesophyll, and is formed of two distinct series of cells, each containing the green chlorophyll-granules, but differing in form and arrangement. Below the epidermis of the upper side of the leaf there are one or two layers of cells, elongated at right angles to the leaf surface (fig. 2, ps), and applied so closely to each other as to leave From Strasburger's Lehrbuch der Botanik by permission of Gustav Fischer.

FIG. 1. - Apex of a shoot showing origin of leaves: f, leaf rudiment; g, rudiment of an axillary bud.

FIG. 2. Section of a Melon leaf, perpendicular to the surface.

Upper epidermis.

Lower epidermis.

Hairs.

Stomata.

Upper (palisade) layers of parenchymatous cells.

Lower (spongy) layers of parenchymatous cells.

Air-spaces connected with stomata. Air-spaces between the loose cells in the spongy parenchyma.

Bundles of fibro-vascular tissue.

Palisade Tissue and Spongy Cells

The upper epidermis alone possesses only small intercellular spaces, except where stomata happen to be present (fig. 2, m); they form the palisade tissue. On the other side of the leaf the cells are irregular, often branched, and are arranged more or less horizontally (fig. 2, p), leaving air-spaces between them, which communicate with stomata; on this account the tissue has received the name of spongy. In leaves having a very firm texture, as those of Coniferae and Cycadaceae, the cells of the parenchyma immediately beneath the epidermis are very much thickened and elongated in a direction parallel to the surface of the leaf, so as to be fibre-like. These constitute a hypodermal layer, beneath which the chlorophyll cells of the parenchyma are densely packed together, and are elongated in a direction vertical to the surface of the leaf, forming the palisade tissue.

Cell Formations

The form and arrangement of the cells, however, depend much on the nature of the plant, and its exposure to light and air. Sometimes the arrangement of the cells on both sides of the leaf is similar, as occurs in leaves which have their edges presented to the sky. In very succulent plants the cells form a compact mass, and those in the centre are often colourless. In some cases the cellular tissue is deficient at certain points, giving rise to distinct holes in the leaf, as in Monstera Adansonii. The fibro-vascular system in the leaf constitutes the venation. The fibro-vascular bundles from the stem bend out into the leaf, and are there arranged in a definite manner. In skeleton leaves, or leaves in which the parenchyma is removed, this arrangement is well seen. In some leaves, as in the barberry, the veins are hardened, producing spines without any parenchyma. The hardening of the extremities of the fibro-vascular tissue is the cause of the spiny margin of many leaves, such as the holly, of the sharp-pointed leaves of madder, and of mucronate leaves, or those having a blunt end with a hard projection in the centre.

Blade/Lamina and Stalk/Petiole

A leaf, whether aerial or submerged, generally consists of a flat expanded portion, called the blade, or lamina, of a narrower portion called the petiole or stalk, and sometimes of a portion at the base of the petiole, which forms a sheath or vagina (fig. 5, s), or is developed in the form of outgrowths, called stipules (fig. 24, s). All these portions are not always present. The sheathing or stipulary portion is frequently wanting. When a leaf has a distinct stalk it is petiolate; when it has none, it is sessile, and if in this case it embraces the stem it is said to be amplexicaul. The part of the leaf next the petiole or the axis is the base, while the opposite extremity is the apex. The leaf is usually flattened and expanded horizontally, i.e. at right angles to the longitudinal axis of the shoot, so that the upper face is directed towards the heavens, and the lower towards the earth. In some cases leaves, as in Iris, or leaf-like petioles, as in Australian acacias and eucalypti, have their plane of expansion parallel to the axis of the shoot, there is then no distinction into an upper and a lower face, but the two sides are developed alike; or the leaf may have a cylindrical or polyhedral form, as in mesembryanthemum. The upper angle formed between the leaf and the stem is called its axil; it is there that leaf-buds are normally developed. The leaf is sometimes articulated with the stem, and when it falls off a scar remains; at other times it is continuous with it, and then decays, while still attached to the axis.

In their early state all leaves are continuous with the stem, and it is only in their after growth that articulations are formed. When leaves fall off annually they are called deciduous; when they remain for two or more years they are persistent, and the plant is evergreen. The laminar portion of a leaf is occasionally articulated with the petiole, as in the orange, and a joint at times exists between the vaginal or stipulary portion and the petiole.

Venation

The arrangement of the fibro-vascular system in the lamina constitutes the venation or nervation. In an ordinary leaf, as that of the elm, there is observed a large central vein running from the base to the apex of the leaf, this is the midrib (fig. 3); it gives off veins laterally (primary veins). A leaf with FIG. 4. - Multicostate leaf of Castoroil plant (Ricinus communis). It is palmately-cleft, and exhibits seven' lobes at the margin. The petiole is inserted a little above the base, and hence the leaf is called peltate or shieldlike.

A leaf with only a single midrib is said to be unicostate and the venation is described as pinnate or feather-veined. In some cases, as sycamore or castor oil (fig. 4), in place of there being only a single midrib there are several large veins (ribs) of nearly equal size, which diverge from the point where the blade joins the petiole or stem, giving off lateral veins. The leaf in this case is multicostate and the venation palmate. The primary veins give off secondary veins, and these in their turn give off tertiary veins, and so on until a complete network of vessels is produced, and those veins usually project on the under surface of the leaf. To a distribution of veins such as this the name of reticulated or netted venation has been applied. In the leaves of some plants there exists a midrib with large veins running nearly parallel to it from the base to the apex of the lamina, as in grasses (fig. 5); or with veins diverging from the base of the lamina in more or less FIG. 3. - Leaf of Elm (Ulmus). (Reticulated venation; primary veins going to the margin, which is serrated. Leaf unequal at the base.)

Some leaves possess veins in parallel lines, as in fan palms (fig. 6), or with veins coming off from it throughout its whole course, and running parallel to each other in a straight or curved direction towards the margin of the leaf, as in plantain and banana. In these cases the veins are often united by cross veinlets, which do not, however, form an angular network. Such leaves are said to be parallel-veined. The leaves of Monocotyledons have generally this kind of venation, while reticulated venation most usually occurs amongst Dicotyledons. Some plants, which in most points of their structure are monocotyledonous, yet have reticulated venation; as in Smilax and Dioscorea. In vascular acotyledonous plants there is frequently a tendency to fork exhibited by the fibro-vascular bundles in the leaf; and when this is the case we have f ork-veined leaves. This is well seen in many ferns. The distribution of the system of vessels in the leaf is FIG. 5. - Stem of a Grass FIG. 6. - Leaf of a Fan Palm (Poa) with leaf. The sheaths (Chamaerops), showing the veins ending in a process 1, called running from the base to the mara ligule; the blade of the gin, and not forming an angular leaf, f. network.

Most veins are usually easily traced, but in the case of succulent plants, as Hoya, agave, stonecrop and mesembryanthemum, the veins are obscure. The function of the veins which consist of vessels and fibres is to form a rigid framework for the leaf and to conduct liquids.

Functions of a Leaf

The form and arrangement of the parts of a typical foliage leaf are intimately associated with the part played by the leaf in the life of the plant. The flat surface is spread to allow the maximum amount of sunlight to fall upon it, as it is by the absorption of energy from the sun's rays by means of the chlorophyll contained in the cells of the leaf that the building up of plant food is rendered possible; this process is known as photo-synthesis; the first stage is the combination of carbon dioxide, absorbed from the air taken in through the stomata into the living cells of the leaf, with water which is brought into the leaf by the wood-vessels. The wood-vessels form part of the fibro-vascular bundles or veins of the leaf and are continuous throughout the leaf-stalk and stem with the root by which water is absorbed from the soil. The palisade layers of the mesophyll contain the larger number of chlorophyll grains (or corpuscles) while the absorption of carbon dioxide is carried on chiefly through the lower epidermis which is generally much richer in stomata. The water taken up by the root from the soil contains nitrogenous and mineral salts which combine with the first product of photo-synthesis - a carbohydrate - to form more complicated nitrogen-containing food substances of a proteid nature; these are then distributed by other elements of the vascular bundles (the phloem) through the leaf to the stem and so throughout the plant to wherever growth or development is going on. A large proportion of the water which ascends to the leaf acts merely as a carrier for the other raw food materials and is got rid of from the leaf in the form of water vapour through the stomata - this process is known as transpiration. Hence the extended surface of the leaf exposing a large area to light and air is eminently adapted for the carrying out of the process of photo-synthesis and transpiration. The arrangement of the leaves on the stem and branches (see Phyllotaxy, below) is such as to prevent the upper leaves shading the lower, and the shape of the leaf serves towards the same end - the disposition of leaves on a branch or stem is often seen to form a "mosaic," each leaf fitting into the space between neighbouring leaves and the branch on which they are borne without overlapping.

Submerged (Underwater) Leaves Structure

Submerged leaves, or leaves which are developed under water, differ in structure from aerial leaves. They have usually no fibro-vascular system, but consist of a congeries of cells, which sometimes become elongated and compressed so as to resemble veins. They have a layer of compact cells on their surface, but no true epidermis, and no stomata. Their internal structure consists of cells, disposed irregularly, and sometimes leaving spaces which are filled with air for the purpose of floating the leaf. When exposed to the air these leaves easily part with their moisture, and become shrivelled and dry. In some cases there is only a network of filament-like cells, the spaces between which are not filled with parenchyma, giving a skeleton appearance to the leaf, as in Ouvirandra fenestralis (Lattice plant).

Simple and Compound Leaf Forms

In all plants, except Thallophytes, leaves are present at some period of their existence. In Cuscuta (Dodder) (q.v.), however, we have an exception. The forms assumed by leaves vary much, not only in different plants, but in the same plant. It is only amongst the lower classes of plants - Mosses, Characeae, &c. - that all the leaves on a plant are similar. As we pass up the scale of plant life we find them becoming more and more variable. The structures in ordinary language designated as leaves are considered so par excellence, and they are frequently spoken of as foliage leaves. In relation to their production on the stem we may observe that when they are small they are always produced in great number, and as they increase in size their number diminishes correspondingly. The cellular process from the axis which develops into a leaf is simple and undivided; it rarely remains so, but in progress of growth becomes segmented in various ways, either longitudinally or laterally, or in both ways. By longitudinal segmentation we have a leaf formed consisting of sheath, stalk and blade; or one or other of these may be absent, and thus stalked, sessile, sheathing, &c., leaves are produced. Lateral segmentation affects the lamina, producing indentations, lobings or fissuring of its margins. In this way two marked forms of leaf are produced - (I) Simple form, in which the segmentation, however deeply it extends into the lamina, does not separate portions of the lamina which become articulated with the midrib or petiole; and (2) Compound form, where portions of the lamina are separated as detached leaflets, which become articulated with the midrib or petiole. In both simple and compound leaves, according to the amount of segmentation and the mode of development of the parenchyma and direction of the fibro-vascular bundles, many forms are produced.

Simple Leaves

When the parenchyma is developed symmetrically on each side of the midrib or stalk, the leaf is equal; if otherwise, the leaf is unequal or oblique (fig. 3). If the margins are even and present no divisions, the leaf is entire (fig. 7); if there are slight projections which are more or less pointed, the leaf is dentate or toothed; when the projections lie regularly over each other, like the teeth of a saw, the leaf is serrate (fig. 3); when they are rounded the leaf is crenate. If the divisions extend more deeply into the lamina than the margin, the leaf receives different names according to the nature of the segments; thus, when the divisions extend about half-way down (fig. 8), it is cleft; when the divisions extend nearly to the base or to the midrib the leaf is partite. If these divisions take place in a simple feather-veined leaf it becomes either pinnatifid (fig. 9), when the segments extend to about the middle, or pinnatipartite, when the divisions extend nearly to the midrib. These primary divisions may be again subdivided in a similar manner, and thus a feather-veined leaf will become bipinnatifid or bipinnatipartite; still further subdivisions give origin to tripinnatifid and laciniated leaves. The same kinds of divisions FIG. 7, ' 'FIG. S.

FIG. 7. - Ovate acute leaf of Coriara myrtifolia. Besides the midrib there are two intra-marginal ribs which converge to the apex. The leaf is therefore tricostate.

FIG. 8. - Runcinate leaf of Dandelion. It is a pinnatifid leaf, with the divisions pointing towards the petiole and a large triangular apex.

FIG. 9. - Pinnatifid leaf of Valeriana dioica. taking place in a simple leaf with palmate or radiating venation, give origin to lobed, cleft and partite forms.

The name palmate or palmatifid (fig. 4) is the general term applied to leaves with radiating venation, in which there are several lobes united by a broad expansioi. of parenchyma, like the palm of the hand, as in the sycamore, castoroil plant, &c. The divisions of leaves with radiating venation may extend to near the base of the leaf, and the names bipartite, tripartite, quinquepartite, &c., are given according as the partitions are two, three, five or more. The term dissected is applied to leaves with radiating venation, having numerous narrow divisions, as in Geranium dissectum. When in a radiating leaf there are three primary partitions, and the two lateral lobes are again cleft, as in hellebore (fig. 11), the leaf is called pedate or pedatifid, from a fancied resemblance to the foot. FIG. 11 - Pedate leaf of Stinking Hellebore (Helleborus foetidus). The venation is radiating. It is a palmately-partite leaf, in which the lateral lobes are deeply divided. When the leaf hangs down it resembles the foot of a bird, and hence the name.

In all the instances already alluded to the leaves have been considered as flat expansions, in which the ribs or veins spread out on the same plane with the stalk. In some cases, however, the veins spread at right angles to the stalk, forming a peltate leaf as in Indian cress (fig. 12).

The form of the leaf shows a very great variety ranging from the narrow linear form with parallel sides, as in grasses or the needle-like leaves of pines and firs to more or less rounded or orbicular - descriptions of these will be found in works on descriptive botany - FIG. 10. - Five-partite leaf of Aconite.


A few examples are illustrated here (figs. 7, 13, 14, 15). The apex also varies considerably, being rounded, or obtuse, sharp or acute (fig. 7), notched (fig. 15), &c. Similarly the shape of the base may vary, when rounded lobes are formed, as in dog-violet, the leaf is cordate or heart-shaped; or kidney-shaped or reniform (fig. 16), when the apex is rounded as in ground ivy. When the lobes are prolonged downwards and are acute, the leaf is sagittate (fig. 17); when they proceed at right angles, as in Rumex Acetosella, the leaf is hastate or halbertshaped. When a simple leaf is divided at the base into two leaf-like appendages, it is called auriculate. When the development of parenchyma is such that it more than fills up the spaces between the veins, the margins become wavy, crisp or undulated, as in Rumex crispus and Rheum undulatum. By cultivation the cellular tissue is often much increased, giving rise to the curled leaves of greens, savoys, cresses, lettuce, &c.

Compound Leaves

Compound leaves are those in which the divisions extend to the midrib or petiole, and the sepa rated portions become each arti culated with it, and receive the name of leaflets. The midrib, or petiole, has thus the appearance of a branch with FIG. 13. - Lanceolate FIG. 12. - Peltate leaves of Indian Cress leaf of a species of (Tropaeolum majus). Senna has separate leaves attached to it, but it is considered properly as one leaf, because in its earliest state it arises from the axis as a single piece, and its subsequent divisions in the form of leaflets are all in one plane. The leaflets are either sessile (fig. 18) or have stalks, called petiolules (fig. 19). Compound leaves are pinnate (fig. 19) or palmate (fig. 18) according to the arrangement of leaflets. When a pinnate leaf ends in a pair of pinnae it is equally or abruptly pinnate (paripinnate); when there is a single terminal leaflet (fig. 19), the leaf is unequally pinnate (imparipinnate); when the leaflets or pinnae are placed alternately on either side of the midrib, and not directly opposite to each other, the leaf is alternately pinnate; and when the pinnae are of different sizes, the leaf is interruptedly pinnate.

FIG. 14. - Oblong leaf of a species of Senna.

FIG. 15. - Emarginate leaf of a species of Senna. The leaf in its contour is somewhat obovate, or inversely egg-shaped, and its base is oblique. FIG. 16. - Reniform leaf of. Nepeta Glechoma, margin crenate. FIG. 17. - Sagittate leaf of Convolvulus.

When the division is carried into the second degree, and the pinnae of a compound leaf are themselves pinnately compound, a bipinnate leaf is formed.

The petiole or leaf-stalk is the part which unites the limb or blade of the leaf to the stem. It is absent in sessile leaves, and this is also C. frequently the case when a sheath is present, as in grasses (fig. 5). It consists of the fibro-vascular bundles with a varying amount of cellular tissue. When the vascular bundles reach the base of the lamina they separate and spread out in various ways, as already described under venation. The lower part of the petiole is often swollen (fig. 20, p), forming the pulvinus, formed of cellular tissue, the cells of which exhibit the phenomenon of irritability. In Mimosa pudica (fig. 20) a sensitiveness is located in the pulvinus which upon irritation induces a depression of the whole bipinnate leaf, a similar property exists in the pulvini at the base of the leaflets which fold upwards. The petiole varies in length, being usually shorter than the lamina, but sometimes much longer. In some palms it is 15 or 20 ft. long, and is so firm as to be used for poles or walking-sticks. In general, the petiole is more or less rounded in its form, the upper surface being flattened or grooved. Sometimes it is compressed laterally, as in the aspen, and to this peculiarity the trembling of the leaves of this tree is due. In aquatic plants the leafstalk is sometimes distended with air, as in Pontederia and Trapa, so as to float the leaf. At other times it is winged, and is either leafy, as in the orange (fig. 21, p), lemon and Dionaea, or pitcherlike, as in Sarracenia (fig. 22). In some Australian acacias, and in some species of Oxalis and Bupleurum, the petiole is flattened in a vertical direction, the vascular bundles separating immediately after quitting the stem and running nearly parallel from base to apex. This kind of petiole (fig. 23, p) has been called a phyllode. In these plants the laminae or blades of the leaves are pinnate or bipinnate and are produced at the FIG. 19. - Imparipinnate (unequally pinnate) leaf of Robinia. There are nine pairs of shortly-stalked leaflets (foliola, pinnae), and an odd one at the extremity. At the base of the leaf the spiny stipules are seen.

Some plants appear with extremities of the phyllodes in a horizontal direction; but in many instances they are not developed, and the phyllode serves the purpose of a leaf. These phyllodes, by their vertical position and their peculiar form, give a remarkable aspect to vegetation. On the same acacia there occur leaves with the petiole and lamina perfect; others having the petiole slightly expanded or winged, and the lamina imperfectly developed; and others in which there is no lamina, and the petiole becomes large and broad. Some petioles are long, slender and sensitive to contact, and function as tendrils by means of which the plant climbs; as in the l,' nasturtiums (Tropaeolum), clematis and c in others; and in compound leaves the midrib and some of the leaflets may similarly be transformed into tendrils, as in the pea and vetch.

The leaf base is of ten developed as a sheath (vagina), which embraces the whole or part of the circumference of the stem (fig. 5). This sheath is comparatively rare in dicotyledons, but is seen in umbelliferous plants. It is much more common amongst monocotyledons. In sedges the sheath forms a complete investment of the stem, whilst in Leaf grasses it is split on one side. In the latter plants there is also a membranous outgrowth, the ligule, at right angles to the median plane of the leaf from the point where the sheath passes into the lamina, there being no petiole (fig. 5, 1). In leaves in which no sheath is produced we not infrequently find small foliar organs, stipules, at the base of the petiole (fig. 24, s). The stipules are generally two in number, and they are important as supplying characters in certain natural orders. Thus they occur FIG. 17.

FIG. 18. - Palmately compound leaf of the Horse-chestnut (Aesculus Hippocastanum). FIG. 20. - Branch and leaves of the Sensitive plant (Mimosa pudica), showing the petiole in its erect state, a, and in its depressed state, b; also the leaflets closed, c, and the leaflets expanded, d.

Irritability resides in the pulvinus,in the pea and bean family, in rosaceous plants and the family Rubiaceae. They are not common in dicotyledons with opposite leaves. Plants having stipules are called stipulate; those having none are exstipulate. Stipules may be large or small, entire or divided, deciduous or persistent. They are not usually of the same form as the ordinary foliage leaves of the plant, from which they are distinguished by their lateral position at the base of the petiole. In the pansy (fig. 24) the true leaves are stalked and crenate, while the stipules s are large, sessile and pin natifid. In Lathyrus Aphaca and some other plants the true pinnate leaves are abortive, the petiole forms a tendril, and the stipules alone are developed, perform ing the office of leaves. When stiP 1? pulate leaves are op posite to each other, at the same height on the stem, it occa FIG. 21. - Leaf of FIG. 22. - Pitcher sionally happens Orange (CitrusAuran- (ascidium) of a species that the stipules on tium), showing a of Side-saddle plant the two sides unite winged leafy petiole p, (Sarracenia purpurea). wholly or partially, which is articulated.

The pitcher is formed so as to form an in- to the lamina 1. from the petiole, which terpetiolary or inter- is prolonged. foliar stipule, as in members of the family Rubiaceae. In the case of alternate leaves, the stipules at the base of each leaf are sometimes united to the petiole and to each other, so as to form an adnate, adherent or petiolary stipule, as in the rose, or an axillary stipule, as in Houttuynia cordata. In other instances the stipules unite together on the side of the stem opposite the leaf forming an ocrea, as in the dock family (fig. 25).

In the development of the leaf the stipules frequently play a most important part. They begin to be formed after the origin of the leaves, but grow much more rapidly than the leaves, and in this way they arch over the young leaves and form protective chambers wherein the parts of the leaf may develop. In the figs, magnolia and pondweeds they are very large and completely envelop the young leaf-bud. The stipules are sometimes so minute as to be scarcely distinguishable without the aid of a lens, and so fugacious as to be visible only in the very young state of the leaf. They may assume a hard and spiny character, as in Robinia Pseudacacia (fig. 19), or may be cirrose, as in Smilax, where each stipule is represented by a tendril. At the base of the leaflets of a compound leaf, small stipules (stipels) are occasionally produced.

Variations in Leaves

Variations in the structure and forms of leaves and leafstalks are produced by the increased development of cellular tissue, by the abortion or degeneration of parts, by the multiplication or repetition of parts and by adhesion. When cellular tissue is developed to a great extent, leaves become succulent and occasionally assume a crisp or curled appearance. Such changes take place naturally, but they are often increased by the art of the gardener, and the object of many horticultural operations is to increase the bulk and succulence of leaves. It is in this way that cabbages and savoys are rendered more delicate and nutritious. By a deficiency in development of parenchyma and an increase in the mechanical tissue, leaves are liable to become hardened and spinescent. The leaves of barberry and of some species of Astragalus, and the stipules of the false acacia (Robinia) are spiny. To the same cause is due the spiny margin of the holly-leaf. When two lobes at the base of a leaf are prolonged beyond the stem and unite (fig. 26), the leaf is perfoliate, the stem appearing to pass through it, as in Bupleurum perfoliatum and Chlora perfoliata; when two leaves unite by their bases they become connate (fig. 27), as in Lonicera Caprifolium; and when leaves adhere to the stem, forming a sort of winged or leafy appendage, they are decurrent, as in thistles. The formation of peltate leaves has been traced to the union of the lobes of a cleft leaf. In the leaf of the Victoria regia the transformation may be traced during germination. The first leaves produced by the young plant are linear,the second are sagittate and hastate, the third are rounded-cordate and the next are orbicular. The stigonum indicating the union of the lobes remains in the Pansy.

Tendrils

The parts of the leaf are frequently transformed into tendrils, with the view of enabling the plants to twine round others for support. In Leguminous plants (the pea tribe) the pinnae are frequently modified to form tendrils, as in Lathyrus Aphaca, in which the stipules perform the function of true leaves. In Flagellaria indica, Gloriosa superba the two lobes at the base of the leaf are united, so that the stalk appears to come through the leaf. and others, the midrib of the leaf ends in a tendril. In Smilax, there are two stipulary tendrils.

Fistular Leaves

The vascular bundles and cellular tissue are sometimes developed in such a way as to form a circle, with a hollow in the centre, and thus give rise to what are called fistular or hollow leaves, as in the onion, and to ascidia or pitchers. Pitchers are formed either by petioles or by laminae, and they are composed of one or more leaves. In Sarracenia (fig. 22) and Heliamphora the pitcher is composed of the petiole of the leaf. In the pitcher plant, Nepenthes, the pitcher is a modification of the lamina, the petiole often plays the part of a tendril, while the leaf base is flat and leaf-like (fig. 28).

In Utricularia bladder-like sacs are formed by a modification of leaflets on the submerged leaves.

In some cases the leaves are reduced to mere scales - cataphyllary leaves; they are produced abundantly upon underground shoots. In parasites (Lathraea, Orobanche) and in plants growing on decaying vegetable matter (saprophytes), in which no chlorophyll is formed, these scales are the only leaves produced. In Pinus the only leaves produced on the main stern and the lateral shoots are scales, the acicular leaves of the tree growing from axillary shoots. In Cycas whorls of scales alternate with large pinnate leaves. In many plants, as already noticed, phyllodia or stipules perform the function of leaves. The production of leaf-buds from FIG. 23. - Leaf of an Acacia (Acacia heterophylla), showing a flattened leaf-like petiole p, called a phyllode, with straight venation, and a bipinnate lamina.

FIG. 27. - Connate leaves of a species of Honeysuckle (Lonicera Caprifolium). Two leaves are united by their bases.

FIG. 28. - Pitcher of a species of pitcher-plant (Nepenthes distillatoria). leaves sometimes occurs as in Bryophyllunt, and many plants of the order Gesneraceae. The leaf of Venus's fly-trap (Dionaea muscipula) when cut off and placed in damp moss, with a pan of water underneath and a bell-glass for a cover, has produced buds from which young plants were obtained. Some species of saxifrage and of ferns also produce buds on their leaves and fronds. In Nymphaea micrantha buds appear at the upper part of the petiole.

Leaf Positioning

Leaves occupy various positions on the stem and branches, and have received different names according to their situation.

Thus leaves arising from the crown of the root, as in Phyllo- the primrose, are called radical; those on the stem are cauline; on flower-stalks, floral leaves (see Flower). The first leaves developed are known as seed leaves or cotyledons. The arrangement of the leaves on the axis and its appendages is called phyllotaxis. In their arrangement leaves follow a definite order. The points on the stem at which leaves appear are called nodes; the part of the stem between the nodes is the internode. When two leaves are produced at the same node, one on each side of the stem or axis, and at the same level, they are opposite (fig. 29); when more than two are produced they are verticillate, and the circle of leaves is then called a verticil or whorl. When leaves are opposite, each successive pair may be placed at right angles to the pair immediately preceding.

They are then said to decussate, following thus a law of alternation (fig. 29). The same occurs in the verticillate arrangement, the leaves of each whorl rarely being super- posed on those of the whorl next it, but usually alterna ting so that each leaf in a whorl occupies the space be tween two leaves of the whorl next to it. There are con siderable irregularities, how ever, in this respect, and the number of leaves in different whorls is not always uniform, as may be seen in Lysimachia

FIG. 29. - A stem FIG. 30. - A vulgaris.

When a single leaf with opposite stem with alteris produced at a node, and leaves. The pairs alternate leaves, and the nodes are separated so are placed at right ranged in a penthat each leaf is placed at angles alternately, tastichous or different height on the stem, or in what is called quincuncial manthe leaves are alternate

In the lowest leaf is directly the point of insertion of the pair one leaf is in above the first, leaf in the node, dividing front and the other and commences the leaf into similar halves, at the back; in the the second cycle. is the median plane of the second pair the The fraction of leaf; and when the leaves are leaves are placed the circumference arranged alternately on an laterally, and so of the stem exaxis so that their median on. pressing the diplanes coincide they form a vergence of the straight row or orthostichy. On every axis there are usually fifths. two or more orthostichies. In fig. 31, leaf 1 arises from a node n; leaf 2 is separated from it by an internode m, and is placed to the right or left; while leaf 3 is situated directly above leaf 1. In this case, then, there are two orthostichies, and the arrangement is said to be distichous. When the fourth leaf is directly above the first, the arrangement is tristichous. The same arrangement continues throughout the branch, so that in the latter case the 7th leaf is above the 4th, the 10th above the 7th; also the 5th above the 2nd, the 6th above the 3rd and so on. The size of the angle between the median planes of two consecutive leaves in an alternate arrangement is their divergence; and it is expressed in fractions of the circumference of the axis which is supposed to be a circle. In a regularlyformed straight branch covered with leaves, if a thread is passed from one to the other, turning always in the same direction, a spiral is described, and a certain number of leaves and of complete turns occur before reaching the leaf directly above that from which the enumeration commenced. If this arrangement is expressed by a fraction, the numerator of which indicates the number of turns, and the denominator the number of internodes in the spiral cycle, the fraction will be found to represent the angle of divergence of the consecutive leaves on the axis. Thus, in fig. 32, a, b, the cycle consists of five leaves, the 6th leaf being placed vertically over the 1st, the 7th over the 2nd and so on; while the number of turns between the 1st and 6th leaf is two; hence this arrangement is indicated by the fraction g. In other words, the distance or divergence between the first and second leaf, expressed in parts of a circle, is of a circle or 360° In fig. 31, a, b, the spiral is 4, i.e. one turn and two leaves; the third leaf being placed vertically over the first, and the divergence between the first and second leaf being one-half the circumference of a circle, 360/2==180°. Again, in a tristichous arrangement the number is or one turn and three leaves, the angular divergence being 120°.

By this means we have a convenient mode of expressing on paper the exact position of the leaves upon an axis. And in many cases such a mode of expression is of excellent service in enabling us readily to understand the relations of the leaves. The divergences may also be represented diagrammatically on a horizontal projection of the vertical axis, as in fig. 33. Here the outermost circle represents a section of that portion of the axis bearing the lowest leaf, the innermost represents the highest. The broad dark lines represent the leaves, and they are numbered according to their age and position. It will be seen at once that the leaves are arranged in orthostichies marked I.-V., and that these divide the circumference into five equal portions. But the divergence between leaf and leaf 2 is equal to tths of the circumference, and the same is the case between 2 and 3, 3 and 4, &c. The divergence, then, is and from this we learn that, starting from any leaf on the axis, we must pass twice round the stem in a spiral through five leaves before reaching one directly over that with which we started. The; line which, winding round an axis either to the right or to the left, passes through the points of insertion of all the leaves on the axis is termed the genetic or generating spiral; and that margin of each leaf which is towards the direction from which the spiral proceeds is the kathodic side, the other margin facing the point whither the spiral passes being the anodic side.

In cases where the internodes are very short and the leaves are closely applied to each other, as in the house-leek, it is difficult to trace the generating spiral. Thus, in fig. 34 there are thirteen leaves which are numbered in their order, and five turns of the spiral marked by circles in the centre (1.5 g indicating the arrangement); but this could not be detected at once. So also in fir cones (fig. 35), which are composed of scales or modified leaves, the generating spiral cannot be determined easily. But in such cases a series of secondary spirals or parastichies are seen running parallel with each other both right and left, which to a certain FIG. 32. - Part of a branch of a extent conceal the genetic spiral. Cherry with six leaves, the sixth The spiral is not always conbeing placed vertically over the stant throughout the whole first, after two turns of the spiral. length of an axis. The angle of This is expressed by two-fifths. divergence may alter either a, The branch, with the leaves abruptly or gradually, and the numbered in order; b, a magnified phyllotaxis thus becomes very representation of the branch, complicated.

This change showing the points of insertion can brought about by arrest of the leaves and their spiral development, by increased development of individual parts or by a torsion of the axis. The former are exemplified in many Crassulaceae and aloes. The latter is seen well in the screw-pine (Pandanus). In the bud of the screw-pine the leaves are arranged in three orthostichies with the phyllotaxis but by torsion the developed leaves become arranged in three strong spiral rows running round the stem. These causes of change in phyllotaxis are also well exemplified in the alteration of an opposite or verticillate arrangement to an alternate, and vice versa; thus the effect of interruption of growth, in causing alternate leaves to become opposite and verticillate, can be distinctly shown in Rhododendron ponticum.

FIG. 31. - Portion of a branch of a Lime tree,with four leaves arranged in a distichous manner, or in two rows. a, The branch with the leaves numbered in their order, n being the node and m the internode; b is a magnified representation of the branch, showing the points of insertion of the leaves and their spiral arrangement, which is expressed by the fraction or one turn of the spiral for two internodes.

The primitive or generating spiral may pass either from right to left or from left to right. It sometimes follows a different direction in the branches from that pursued in the stem. When it follows the same course in the stem and branches, they are homodromous; when the direction differs, they are heterodromous. In different species of the same genus the phyllotaxis frequently varies.

All modifications of leaves follow the same laws of arrangement as true leaves - a fact which is of importance in a morphological point of view. In dicotyledonous plants the first leaves produced (the cotyledons) are opposite. This arrangement often continues during the life of the plant, but at other times it changes, passing into distichous and spiral forms. Some tribes of plants are distinguished by their opposite or ver ticillate, others by their alternate, leaves. Labiate plants have decussate leaves, while Boragin aceae have alternate leaves, and Tiliaceae usua ally have distichous leaves; Rubiaceae have opposite leaves. Such arrangements are common in Dicotyledons. The first of these, called a quincunx, is met with in the apple, pear and cherry (fig. 32); the second, in the bay, holly, Plantago media; the third, in the cones of Picea alba (fig. FIG. 33. - Diagram of a phyllotaxis); and the fourth in those of the silver fir.

Seed Leaves

In monocotyledonous plants there is only one seed-leaf or cotyledon, and hence the arrangement is at first alternate; and it generally continues so more or less, rarely being verticillate. Such arrangements as 2, 3 ands are common in Monocotyledons, as in grasses, sedges and lilies. It has been found in general that, while the number 5 occurs in the phyllotaxis of Dicotyledons, 3 is common in that of Monocotyledons.

In the axil of previously formed leaves leaf-buds arise. These leaf-buds contain the rudiments of a shoot, and consist of leaves covering a growing point. The buds of trees of temperate climates, which lie dormant during the winter, are protected by scale leaves. These scales or protective appendages of the bud consist either of FIG. 34. - Cycle of thirteen leaves placed closely together so as to form FIG. 35. - Cone of Picea alba a rosette, as in Sempervivum. A is with the scales or modified the very short axis to which the leaves numbered in the order leaves are attached. The leaves are of their arrangement on the numbered in their order, from below axis of the cone. The lines upwards. The circles in the centre indicate a rectilinear series of indicate the five turns of the spiral, scales and two lateral secondand show the insertion of each of the ary spirals, one turning from leaves.

The divergence is expressed left to right, the other is right to left, in the altered laminae or of the enlarged petiolary sheath, or of stipules, as in the fig and magnolia, or of one or two of these parts combined. These are often of a coarse nature, serving a temporary purpose, and then falling off when the leaf is expanded. They are frequently covered with a resinous matter, as in balsam-poplar and horsechestnut, or by a thick downy covering as in the willow. In plants of warm climates the buds have often no protective appendages, and are then said to be naked.

The arrangement of the leaves in the bud is termed vernation or prefoliation. In considering vernation we must take into account both the manner in which each individual leaf is folded and also the arrangement of the leaves in relation to each other. These vary in different plants, but in each species they follow a regular law. The leaves in the bud are either placed simply in apposition, as in the mistletoe, or they are folded or rolled up longitudinally or laterally, giving rise to different kinds of vernation, as delineated in figs. 36 to 45, where the folded or curved lines represent the leaves, the thickened part being the midrib. The leaf taken individually is either folded longitudinally from apex to base, as in the tulip-tree, and called reclinate or replicate; or rolled up in a circular manner from apex to base, as in ferns (fig. 36), and called circinate; or folded laterally, conduplicate (fig. 37), as in oak; or it has several folds like a fan, plicate or plaited (fig. 38), as in vine and sycamore, and in leaves with radiating vernation, where the ribs mark the foldings; or it is rolled upon itself, convolute (fig. 39), as in banana and apricot; or its edges are rolled inwards, involute (fig. 40), as in violet; or outwards, revolute (fig. 41), as in rosemary. The different divisions of a cut leaf may be folded or rolled up separately, as in ferns, while the entire leaf may have either the same or a different kind of vernation. The leaves have a definite relation to each other in the bud, being either opposite, alternate or verticillate; and thus different kinds of vernation are produced. Sometimes they are nearly in a circle at the same level, remaining flat or only slightly convex externally, and placed so as to touch each other by their edges, thus giving rise to valvate vernation. At other times they are at different levels, and are applied over each other, so as to be imbricated, as in lilac, and in the outer scales of sycamore; and occasionally the margin of one leaf overlaps that of another, while it in its turn is overlapped by a third, so as to be twisted, spiral or contortive.

FIG. 36. - Circinate vernation. FIG. 37. - Transverse section of a conduplicate leaf. FIG. 38. - Transverse section of a plicate or plaited leaf. FIG. 39. - Transverse section of a convolute leaf. FIG. 40. - Transverse section of an involute leaf. FIG. 41. - Transverse section of a revolute leaf.

FIG. 42. - Transverse section of a bud, in which the leaves are arranged in an accumbent manner.

FIG. 43. - Transverse section of a bud, in which the leaves are arranged in an equitant manner.

FIG. 44. - Transverse section of a bud, showing two leaves folded in an obvolute manner. Each is conduplicate, and one embraces the edge of the other.

FIG. 45. - Transverse section of a bud, showing two leaves arranged in a supervolute manner.


When leaves are applied to each other face to face, without being folded or rolled together, they are appressed. When the leaves are more completely folded they either touch at their extremities and are accumbent or opposite (fig. 42), or are folded inwards by their margin and become induplicate; or a conduplicate leaf covers another similarly folded, which in turn covers a third, and thus the vernation is equitant (fig. 43), as in privet; or conduplicate leaves are placed so that the half of the one covers the half of another, and thus they become half-equitant or obvolute (fig. 44), as in sage. When in the case of convolute leaves one leaf is rolled up within the other, it is supervolute (fig. 45). The scales of a bud sometimes exhibit one kind of vernation and the leaves another. The same modes of arrangement occur in the flower-buds.

FIG. 38.

Leaf Death

Leaves, after performing their functions for a certain time, wither and die. In doing so they frequently change colour, and hence arise the beautiful and varied tints of the autumnal foliage. This change of colour is chiefly occasioned by the diminished circulation in the leaves, and the higher degree of oxidation to which their chlorophyll has been submitted.

Leaves which are articulated with the stem, as in the walnut and horse-chestnut, fall and leave a scar, while those which are continuous with it remain attached for some time after they have lost their vitality. Most of the trees of Great Britain have deciduous leaves, their duration not extending over more than a few months, while in trees of warm climates the leaves often remain for two or more years. In tropical countries, however, many trees lose their leaves in the dry season. The period of defoliation varies in different countries according to the nature of their climate. Trees which are called evergreen, as pines and evergreen-oak, are always deprived of a certain number of leaves at intervals, sufficient being left, however, to preserve their green appearance. The cause of the fall of the leaf in cold climates seems to be deficiency of light and heat in winter, which causes a cessation in the functions of the cells of the leaf. The fall is directly caused by the formation of a layer of tissue across the base of the leaf-stalk; the cells of this layer separate from one another and the leaf remains attached only by the fibres of the veins until it becomes finally detached by the wind or frost. Before its fall the leaf has become dry owing to loss of water and the removal of the protoplasm and food substances to the stem for use next season; the red and yellow colouring matters are products of decomposition of the chlorophyll. Inorganic and other waste matters are stored in the leaf-tissue and thus got rid of by the plant. The leaf scar is protected by a corky change (suberization) in the walls of the exposed cells. (A. B. R.)


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Bible wiki

Up to date as of January 23, 2010

From BibleWiki


of a tree. The olive-leaf mentioned Gen. 8:11. The barren fig-tree had nothing but leaves (Matt. 21:19; Mark 11:13). The oak-leaf is mentioned Isa. 1:30; 6:13. There are numerous allusions to leaves, their flourishing, their decay, and their restoration (Lev. 26:36; Isa. 34:4; Jer. 8:13; Dan. 4:12, 14, 21; Mark 11:13; 13:28). The fresh leaf is a symbol of prosperity (Ps. 1:3; Jer. 17:8; Ezek. 47:12); the faded, of decay (Job 13:25; Isa. 1:30; 64:6; Jer. 8:13).

Leaf of a door (1 Kings 6:34), the valve of a folding door.

Leaf of a book (Jer. 36:23), perhaps a fold of a roll.

This entry includes text from Easton's Bible Dictionary, 1897.

what mentions this? (please help by turning references to this page into wiki links)


Gaming

Up to date as of February 01, 2010

From Wikia Gaming, your source for walkthroughs, games, guides, and more!

Leaf is a Japanese visual novel studio under the publisher AQUAPLUS (アクアプラス), and has offices in Yodogawa-ku, Osaka, and Tokyo. It and its competitor Key (to which it is often compared) are the most popular and successful dedicated visual novel studios operating today. It was launched out of obscurity by its early release ToHeart.

Leaf used the XviD video codec in several games: Aruru to Asobo!!, Tears to Tiara, Kusari, and ToHeart2 XRATED. Since XviD is free software, released under the GPL, Leaf was forced to release the source code to those games under the same license. One still required the game data to actually play the games with the source code.[1][2]

A free software engine called xlvns was derived from this source code.[3]

Contents

P/ECE

In addition of software publishing, Aquaplus produced P/ECE, a greyscale mobile gaming platform that allows user to download games via USB or infrared port. The unit has 256KB SRAM and RAM 512KB flash memory.

P/ECE unit has preloaded 'Picket' software, which is an electronic organizer. Aquaplus offered downloadable freeware titles such as Multi's going out, Black Wings, TANK BATTLE, Ojomajomini, Inagawa de urou!!.

Titles

  • DR2 Night Janki (1995)
  • Filsnown (1995)
  • Leaf Visual Novel Series (LVNS)
    • Shizuku (1996)
    • Kizuato (1996)
    • ToHeart (1997)
  • White Album (1998)
  • Comic Party (1999)
  • Magical Antique (2000)
  • Tasogare (2001)
  • Utawarerumono (2002)
  • Routes (2003)
  • December When There Is No Angel (2003)
  • Aruru to Asobo!! (2004)
  • Tears to Tiara (2005)
  • Kusari (2005)
  • ToHeart2 XRATED (2005)
  • FullAni (2006)

References

Smallwikipedialogo.png This page uses content from Wikipedia. The original article was at Leaf (company). The list of authors can be seen in the page history. As with Wikia Gaming, the text of Wikipedia is available under the Creative Commons Attribution-Share Alike 3.0 (unported) license.
  1. http://leaf.aquaplus.co.jp/product/xvid.html
  2. http://ja.wikinews.org/wiki/Leaf、一部作品のソースコード公開へ
  3. http://web.archive.org/web/20031208181123/http://leafbsd.denpa.org/

External links

  • Leaf
  • AquaPlus
  • P/ECE Official WebPage

This article uses material from the "Leaf" article on the Gaming wiki at Wikia and is licensed under the Creative Commons Attribution-Share Alike License.

Simple English

[[File:||thumb|right|200px|The leaves of a Beech tree]]

File:Leaf 1
A leaf in summer. This picture also shows the veins of the leaf.
File:Autumn
Fallen autumn leaves

In botany, a leaf is an above-ground plant organ. Originally, leaves were used for photosynthesis. A leaf is often flat in order to absorb the most light, and thin so that the sunlight will shine through it. Leaves stometimes store food or water. Some plants have changed some of their leaves to do other things.

Those plants that have leaves can be sorted into two groups: Those that have leaves all year round, and those that do not. The plants that do not have leaves all year round generally lose their leaves in autumn. Before this happens, the leaves change colour. The leaves will grow back in spring. Leaves come in many shapes and sizes that help what the certain leaf does for the plant it's on.

Leaves can have different functions

Most of the time, leaves are used to do photosynthesis, they make carbon dioxide and sugar from oxygen using water. Sometimes leaves have been changed to provide a different function:

  • Thorns help protect the plant from being eaten.
  • Vines help the plant to attach to surfaces, and to climb.
  • Some leaves are used to store energy. An example for this is the onion.
  • Many succulents store water in some of their leaves.
  • Some plants (called Epiphytes) grow on other plants. They do not have roots, usually. Often, they changed their leaves to be able to capture rainwater.
  • Carnivorous plants use specially-designed leaves to capture their prey.
File:Leaf morphology no
Leaves can have different shapes. The part of biology that studies the shapes of things is called Morphology

[[File:|thumb|leaves]]








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