Leaf: Encyclopedia II - Leaf - Leaf morphology

Leaf - Leaf morphology

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 continue to grow as long as they have the resources to do so.

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

Leaf - Basic leaf types

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

Leaf - Arrangement on the stem

As a stem grows, leaves tend to appear arranged around the stem in away that optimizes yield of light. In essence, leaves come off the stem in a spiral pattern, 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 series: 1/2, 2/3, 3/5, 5/8, 8/13, 13/21, 21/34, 34/55, 55/89. This series tends to a limit of 360° x 34/89 = 137.52 or 137° 30', an angle known mathematically as the 'golden angle'. In the series, the numerator gives the number of complete turns or gyres until the leaf arrives at the initial position. The denominator gives 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.

The fact that an arrangement of anything in nature can be described by a mathematical formula is not in itself mysterious. Mathematics is the science of discovering numerical relationships and applying formulae to these relationships. The formulae themselves can provide clues to the underlying physiological processes that, in this case, determine where the next leaf bud will form in the elongating stem. However, we can more easily describe the arrangement of leaves using the following terms:

• Alternate — leaf attachments singular at nodes, and leaves alternate direction, to a greater or lesser degree, along the stem.
• Opposite — leaf attachments paired at each node; decussate if, as typical, each successive pair is rotated 90° going along the stem; or distichous if not rotated, but two-ranked (in the same plane).
• Whorled — three or more leaves attach at each point or node on the stem. As with opposite leaves, successive whorls may or may not be decussate, rotated by half the angle between the leaves in the whorl (i.e., successive whorls of three rotated 60°, whorls of four rotated 45°, etc). Note: opposite leaves may appear whorled near the tip of the stem.
• Rosulate — leaves form a rosette ( = a cluster of leaves growing in crowded circles from a common center).

Leaf - Divisions of the lamina blade

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 one 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.

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

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

In some Acacia species, such as the Koa Tree (Acacia koa), the petioles are expanded or broadened and function like leaf blades; these are called phyllodes. There may or may not be normal pinnate leaves at the tip of the phyllode.

• A stipule, present on the leaves of many dicotyledons, is an appendage on each side at the base of the petiole, resembling a small leaf. They may be lasting and not be shed (a stipulate leaf, such as in roses and beans); or be shed as the leaf expands, leaving a stipule scar on the twig (an exstipulate leaf).
• The situation, arrangement, and structure of the stipules is called the stipulation.
  • free
  • adnate : fused to the petiole base
  • ochreate : provided with ochrea, or sheath-formed stipules, e.g. rhubarb,
  • encircling the petiole base
  • interpetiolar : between the petioles of two opposite leaves.
  • intrapetiolar : between the petiole and the subtending stem

Leaf - Venation arrangement of the veins

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

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

Adapted from the Wikipedia article "Leaf morphology", under the G.N U Free Docmentation License. Please also see http://en.wikipedia.org/wiki