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A leaf is an organ of a vascular plant and is the principal lateral appendage of the stem.[1] The leaves and stem together form the shoot.[2] Leaves are collectively referred to as foliage, as in "autumn foliage."[3][4]
Although leaves can be seen in many different textures and sizes, typically a leaf is a thin, dorsiventrally flattened organ, borne above ground and specialized for photosynthesis. In most leaves, the primary photosynthetic tissue, the palisade mesophyll, 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 have distinctive upper surface (adaxial) and lower surface (abaxial) that differ in colour, hairiness, the number of stomata (pores that intake and output gases), epicuticular wax amount and structure and other features.
Broad, flat leaves with complex venation are known as megaphylls and the species that bear them, the majority, as broad-leaved or megaphyllous plants. In others, such as the clubmosses, 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, and 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.[4][7] Many structures of non-vascular plants, such as the phyllids of mosses and liverworts and even of some foliose lichens, which are not plants at all (in the sense of being members of the kingdom Plantae), look and function much like leaves.
- 1 General characteristics
- 2 Morphology (large-scale features)
- 3 Terminology
- 4 Anatomy (medium and small scale)
- 5 Leaf development
- 6 Adaptations
- 7 Seasonal leaf loss
- 8 Interactions with other organisms
- 9 See also
- 10 References
- 11 Bibliography
- 12 External links
General characteristics[edit]
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. Also conifers, whose leaves are needle shaped. The flat, or laminar, shape also maximises thermal contact with the surrounding air. The leaf shape also minimises damage from wind, by creating turbulence rather than resistance.[citation needed] Functionally, in addition to photosynthesis the leaf is the principal site of transpiration and guttation. Leaves also function to store chemical energy and water (especially succulents) and may become specialised organs serving other functions.[which?][8]
The internal organisation of most kinds of leaves has evolved to maximise exposure of the photosynthetic organelles, the chloroplasts, to light and to increase the absorption of carbon dioxide. Their surfaces are waterproofed by the plant cuticle and gas exchange between the mesophyll cells and the atmosphere is controlled by minute openings called stomata, about 10 μm which open or close to regulate the rate exchange of carbon dioxide, oxygen, and water vapour 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.[9]
Many gymnosperms 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 favour 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]
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 neighbours; however, such trees tend to have erect or horizontal dorsiventral leaves as seedlings, when their growth is limited by the available light.[14] Other factors include the need to balance water loss at high temperature and low humidity against the need to absorb atmospheric carbon dioxide. In most plants leaves also are the primary organs responsible for transpiration and guttation (beads of fluid forming at leaf margins).
Leaves can also store food and water, and are modified accordingly to meet these functions, for example in the leaves of succulent plants and in bulb scales. The concentration of photosynthetic structures in leaves requires that they be richer in protein, minerals, and sugars than, say, woody stem tissues. Accordingly, leaves are prominent in the diet of many animals.
Correspondingly, leaves represent heavy investment on the part of the plants bearing them, and their retention or disposition are the subject of elaborate strategies for dealing with pest pressures, seasonal conditions, and protective measures such as the growth of thorns and the production of phytoliths, lignins, tannins and poisons.
Deciduous plants in frigid or cold temperate regions typically shed their leaves in autumn, whereas in areas with a severe dry season, some plants may shed their leaves until the dry season ends. In either case the shed leaves may be expected to contribute their retained nutrients to the soil where they fall.
In contrast, many other non-seasonal plants, such as palms and conifers, retain their leaves for long periods; Welwitschia retains its two main leaves throughout a lifetime that may exceed a thousand years.
The leaf-like organs of Bryophytes (e.g., mosses and liverworts), known as phyllids, differ morphologically from the leaves of vascular plants in that they lack vascular tissue, are usually only a single cell thick and have no cuticle stomata or internal system of intercellular spaces.
Simple, vascularised leaves (microphylls) first evolved as enations, extensions of the stem, in clubmosses such as Baragwanathia during the Silurian period. True leaves or euphylls of larger size and with more complex venation did not become widespread in other groups until the Devonian period, by which time the carbon dioxide concentration in the atmosphere had dropped significantly. This occurred independently in several separate lineages of vascular plants, in progymnosperms like Archaeopteris, in Sphenopsida, ferns and later in the gymnosperms and angiosperms. Euphylls are also referred to as macrophylls or megaphylls (large leaves).[6]
Morphology (large-scale features)[edit]
A structurally complete leaf of an angiosperm consists of a petiole (leaf stalk), a lamina (leaf blade), and stipules (small structures located to either side of the base of the petiole). Not every species produces leaves with all of these 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 morphology. The petiole mechanically links the leaf to the plant and provides the route for transfer of water and sugars to and from the leaf. The lamina is typically the location of the majority of photosynthesis. The upper (adaxial) angle between a leaf and a stem is known as the axil of the leaf. It is often the location of a bud. Structures located there are called "axillary".
External leaf characteristics, such as shape, margin, hairs, the petiole, and the presence of stipules, are important for identifying plant species, and botanists have developed a rich terminology for describing leaf characteristics. Leaves have determinate growth. They grow to a specific pattern and shape and then stop. Other plant parts like stems or roots have non-determinate growth, and will usually continue to grow as long as they have the resources to do so.
The type of leaf is usually characteristic of a species (monomorphic), although some species produce more than one type of leaf (dimorphic or polymorphic). The longest leaves are those of the Raffia palm, R. regalis which may be up to 25 m (82 ft) long and 3 m (9.8 ft) wide.[16] The terminology associated with the description of leaf morphology is presented, in illustrated form, at Wikibooks.
Where leaves are basal, and lie on the ground, they are referred to as prostrate.
Basic leaf types[edit]
- Ferns have fronds
- Conifer leaves are typically needle- or awl-shaped or scale-like
- Angiosperm (flowering plant) leaves: the standard form includes stipules, a petiole, and a lamina
- Lycophytes have microphyll leaves.
- Sheath leaves (type found in most grasses and many other monocots)
- Other specialized leaves (such as those of Nepenthes, a pitcher plant)
Arrangement on the stem[edit]
Different terms are usually used to describe the arrangement of leaves on the stem (phyllotaxis):
- Alternate
- One leaf, branch, or flower part attaches at each point or node on the stem, and leaves alternate direction, to a greater or lesser degree, along the stem.
- Basal
- Arising from the base of the stem.
- Cauline
- Arising from the aerial stem.
- Opposite
- Two leaves, branches, or flower parts attach at each point or node on the stem. Leaf attachments are paired at each node and decussate if, as typical, each successive pair is rotated 90° progressing along the stem.
- Whorled, or verticillate
- Three or more leaves, branches, or flower parts attach at each point or node on the stem. As with opposite leaves, successive whorls may or may not be decussate, rotated by half the angle between the leaves in the whorl (i.e., successive whorls of three rotated 60°, whorls of four rotated 45°, etc.). Opposite leaves may appear whorled near the tip of the stem. Pseudoverticillate describes an arrangement only appearing whorled, but not actually so.
- Rosulate
- Leaves form a rosette.
- Rows
- The term, distichous, literally means two rows. Leaves in this arrangement may be alternate or opposite in their attachment. The term, 2-ranked, is equivalent. The terms, tristichous and tetrastichous, are sometimes encountered. For example, the "leaves" (actually microphylls) of most species of Selaginella are tetrastichous, but not decussate.
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 centered 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 the golden angle, which is approximately 360° × 34/89 ≈ 137.52° ≈ 137° 30′. In the series, the numerator indicates the number of complete turns or "gyres" until a leaf arrives at the initial position and 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): 3 leaves in 1 circle
- 144° (or 2/5): 5 leaves in 2 gyres
- 135° (or 3/8): 8 leaves in 3 gyres.
Divisions of the blade[edit]
Two basic forms of leaves can be described considering the way the blade (lamina) 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 being separated along a main or secondary vein. The leaflets may have petiolules and stipels, the equivalents of the petioles and stipules of leaves. Because each leaflet can appear to be a simple leaf, it is important to recognize where the petiole occurs to identify a compound leaf. Compound leaves are a characteristic of some families of higher plants, such as 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 of 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.
- 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 group of pinnules on each secondary vein forms a "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[edit]
Petiolated leaves have a petiole (leaf stalk), and are said to be petiolate.
Sessile (epetiolate) leaves have no petiole and the blade attaches directly to the stem. Subpetiolate leaves are nearly petiolate or have an extremely short petiole and may appear to be sessile.
In clasping or decurrent leaves, the blade partially surrounds the stem.
When the leaf base completely surrounds the stem, the leaves are said to be perfoliate, such as in Eupatorium perfoliatum.
In peltate leaves, the petiole attaches to the blade inside 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, lateral
- As in Hibiscus.
- Adnate
- Fused to the petiole base, as in Rosa.
- Ochreate
- Provided with ochrea, or sheath-formed stipules, as in Polygonaceae; e.g., rhubarb.
- Encircling the petiole base
- Interpetiolar
- Between the petioles of two opposite leaves, as in Rubiaceae.
- Intrapetiolar
- Between the petiole and the subtending stem, as in Malpighiaceae.
Veins (sometimes referred to as nerves) constitute one of the more visible leaf traits or characteristics. The veins in a leaf represent the vascular structure of the organ, extending into the leaf via the petiole and provide transportation of water and nutrients between leaf and stem, and play a crucial role in the maintenance of leaf water status and photosynthetic capacity.They also play a role in the mechanical support of the leaf.[17][18] Within the lamina of the leaf, while some vascular plants possess only a single vein, in most this vasculature generally divides (ramifies) according to a variety of patterns (venation) and form cylindrical bundles, usually lying in the median plane of the mesophyll, between the two layers of epidermis.[19] This pattern is often specific to taxa, and of which angiosperms possess two main types, parallel and reticulate (net like). In general, parallel venation is typical of monocots, while reticulate is more typical of eudicots and magnoliids ("dicots"), though there are many exceptions.[20][19][21]
The vein or veins entering the leaf from the petiole are called primary or fist order veins. The veins branching from these are secondary or second order veins. These primary and secondary veins are considered major veins or lower order veins, though some authors include third order.[22] Each subsequent branching is sequentially numbered, and these are the higher order veins, each branching being associated with a narrower vein diametre.[23] In parallel veined leaves, the primary veins run parallel and equidistant to each other for most of the length of the leaf and then converge or fuse (anastomose) towards the apex. Usually many smaller minor veins interconnect these primary veins, but may terminate with very fine vein endings in the mesophyll. Minor veins are more typical of angiosperms, which may have as many as four higher orders.[22] In contrast, leaves with reticulate venation there is a single (sometimes more) primary vein in the centre of the leaf, referred to as the midrib or costa and is continuous with the vasculature of the petiole more proximally. The midrib then branches to a number of smaller secondary veins, also known as second order veins, that extend toward the leaf margins. These often terminate in a hydathode, a secretory organ, at the margin. In turn, smaller veins branch from the secondary veins, known as tertiary or third order (or higher order) veins, forming a dense reticulate pattern. The areas or islands of mesophyll lying between the higher order veins, are called areoles. Some of the smallest veins (veinlets) may have their endings in the areoles, a process known as areolation.[23] These minor veins act as the sites of exchange between the mesophyll and the plant's vascular system.[18] Thus minor veins collect the products of photosynthesis (photosynthate) from the cells where it takes place, while major veins are responsible for its transport outside of the leaf. At the same time water is being transported in the opposite direction.[24][20][19]
The number of vein endings is very variable, as is whether second order veins end at the margin, or link back to other veins.[21] There are many elaborate variations on the patterns that the leaf veins form, and these have functional implications. Of these, angiosperms have the greatest diversity.[22] Within these the major veins function as the support and distribution network for leaves and are correlated with leaf shape. For instance the parallel venation found in most monocots correlates with their elongated leaf shape and wide leaf base, while reticulate venation is seen in simple entire leaves, while digitate leaves typically have venation in which three or more primary veins diverge radially from a single point.[25][18][23][26]
In evolutionary terms, early emerging taxa tend to have dichotomous branching with reticulate systems emerging later. Veins appeared in the Permian period (299–252 mya), prior to the appearance of angiosperms in the Triassic (252–201 mya), during which vein hierarchy appeared enabling higher function, larger leaf size and adaption to a wider vaiety of climatic conditions.[22] 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[edit]
- Homoblasty
- Characteristic in which a plant has small changes in leaf size, shape, and growth habit between juvenile and adult stages, in contrast to;
- Heteroblasty
- Characteristic in which a plant has marked changes in leaf size, shape, and growth habit between juvenile and adult stages.
Edge (margin)[edit]
|Even; with a smooth margin; without toothing
|Fringed with hairs
|Wavy-toothed; dentate with rounded teeth
May be coarsely dentate, having large teeth
or glandular dentate, having teeth which bear glands
|Finely toothed
|Doubly serrate
|Each tooth bearing smaller teeth
|Saw-toothed; with asymmetrical teeth pointing forward
|Finely serrate
|With deep, wave-like indentations; coarsely crenate
|Indented, with the indentations not reaching the center
|With a wavy edge, shallower than sinuate
|Spiny or pungent
|With stiff, sharp points such as thistles
- 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.
- Coriaceous
- Leathery; stiff and tough, but somewhat flexible.
- 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.
- Lepidote
- Coated with small scales (thus elepidote, without such scales).
- Maculate
- Stained, spotted, compare immaculate.
- 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.
"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.
- Glabrescent
- Losing hairs with age.
- 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.
- Tomentulose
- Minutely or only slightly tomentose.
- Villous
- With long and soft hairs, usually curved.
- Woolly
- With long, soft and tortuous or matted hairs.
- Hysteranthous
- Developing after the flowers [27]
- Synanthous
- Developing at the same time as the flowers [28]
A number of different classification systems of the patterns of leaf veins (venation) have been described,[21] starting with Ettingshausen (1861),[29] and the terminology has ben described as "formidable".[21] One of the commonest among these is the Hickey system, originally developed for "dicotyledons" and using a number of Ettingshausen's terms derived from Greek (1973–1979):[30][31][32] (see also: Simpson Figure 9.12, p. 468)[21]
- 1. Pinnate (feather-veined, reticulate, pinnate-netted, penniribbed, penninerved, or penniveined)
The veins arise pinnately (feather like) from a single primary vein (mid-vein) and subdivide into secondary veinlets, known as higher order veins. These, in turn, form a complicated network. This type of venation is typical for (but by no means limited to) "dicotyledons" (non monocotyledon angiosperms). E.g. Ostrya There are three subtypes of pinnate venation;
- Craspedodromous (Greek: kraspedon - edge, dromos - running): The major veins reach to the margin of the leaf.
- Camptodromous: Major veins extend close to the margin, but bend before they intersect with the margin.
- Hyphodromous: all secondary veins are absent, rudimentary or concealed
These in turn have a number of further subtypes such as eucamptodromous, where secondary veins curve near the margin without joining adjacent secondary veins.
- 2. Parallelodromous (Parallel-veined, parallel-ribbed, parallel-nerved, penniparallel, striate)
Two or more primary veins originating beside each other at the leaf base, and running parallel to each other to the apex and then converging there. Commissural veins (small veins) connect the major parallel veins. Typical for most monocotyledons, such as grasses. The additional terms marginal (primary veins reach the margin), and reticulate (primary veins do not reach the margin) are also used.
- 3. Campylodromous (campylos - curve)
Several primary veins or branches originating at or close to a single point and running in recurved arches, then converging at apex. E.g. Maianthemum
- 4. Acrodromous
Two or more primary or well developed secondary veins in convergent arches towards apex, without basal recurvature as in Campylodromous. May be basal or suprabasal depending on origin, and perfect or imperfect depending on whether they reach to 2/3 of the way to the apex. E.g. Miconia (basal type), Endlicheria (suprabasal type)
- 5. Actinodromous
Three or more primary veins diverging radially from a single point. E.g. Arcangelisia (basal type), Givotia (suprabasal type)
- 6. Palinactodromous
Primary veins with one or more points of secondary dichotomous branching beyond the primary divergence, either closely or more distantly spaced.E.g. Platanus
Types 4–6 may similarly be subclassified as basal (primaries joined at the base of the blade) or suprabasal (diverging above the blade base), and perfect or imperfect, but also flabellate.
At about the same time, Melville (1976) described a system applicable to all Angiosperms and using Latin and English terminology.[33] Melville also had six divisions, based on the order in which veins develop.
- Arbuscular (arbuscularis) - branching repeatedly by regular dichotomy to give rise to a three dimensional bush-like structure consisting of linear segment (2 subclasses)
- Flabellate (flabellatus) - primary veins straight or only slightly curved, diverging from the base in a fan-like manner (4 subclasses)
- Palmate (palmatus) - curved primary veins (3 subclasses)
- Pinnate (pinnatus) - single primary vein, the midrib, along which straight or arching secondary veins are arranged at more or less regular intervals (6 subclasses)
- Collimate (collimatus) - numerous longitudinally parallel primary veins arising from a transverse meristem (5 subclasses)
- Conglutinate (conglutinatus) - derived from fused pinnate leaflets (3 subclasses)
A modified form of the Hickey system was later incorporated into the Smithsonian classification (1999) which proposed seven main types of venation, based on the architecture of the primary veins, adding Flabellate as an additional main type. Further classification was then made on the basis of secondary veins, with 12 further types, such as;
- brochidodromous (closed form in which the secondaries are joined together in a series of prominent arches, as in Hildegardia)
- craspedodromous (open form with secondaries terminating at the margin, in toothed leaves, as in Celtis)
- eucamptodromous (intermediate form with upturned secondaries that gradually diminish apically but inside the margin, and connected by intermediate tertiary veins rather than loops between secondaries, as in Cornus)
- cladodromous (secondaries freely branching toward the margin, as in Rhus)
terms which had been used as subtypes in the original Hickey system.[34]
Further descriptions included the higher order, or minor veins and the patterns of areoles (see Leaf Architecture Working Group, Figures 28–29).[34]
- Flabellate
Several to many equal fine basal veins diverging radially at low angles and branching apically. E.g. Paranomus
Analyses of vein patterns often fall into consideration of the vein orders, primary vein type, secondary vein type (major veins), and minor vein density. A number of authors have adopted simplified versions of these schemes.[35][21] At its simplest the primary vein types can be considered in three or four groups depending on the plant divisions being considered;
- pinnate
- palmate
- parallel
where palmate refers to multiple primary veins that radiate from the petiole, as opposed to branching from the central main vein in the pinnate form, and encompasses both of Hickey types 4 and 5, which are preserved as subtypes, e.g. palmate-acrodromous (see National Parks Service Leaf Guide).[36]
- Palmate, 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).
Alternatively Simpson uses:[21]
- uninervous - central midrib with no lateral veins (microphyllous), seen in the non-seed bearing tracheophytes, such as horsetails
- dichotomous - veins successively branching into equally sized veins, E.g. Ginkgo and some pteridophytes
- parallel - primary and secondary veins parallel to each other
- netted (reticulate, pinnate) - ultimate veinlets forming an interconnecting net like pattern; most non-monocot angiosperms
However, these simplified systems allow for further division into multiple subtypes. Simpson divides parallel and netted as follows;
- Parallel
- Penni-parallel (pinnate parallel) - secondary veins from a single primary, and parallel, E.g. Zingiberales
- Palmate-parallel - several primary veins from a single point and secondary veins parallel, E.g. "fan" (palmate) palms
- Netted
- Pinnately (veined or netted) - secondary veins along the length of the primary midrib
- Palmately - four or more primary veins arising from a single point
- Ternately - three primary veins, as above
These complex systems are not used much in morphological descriptions of taxa, but have usefulness in plant identification. [21]
Other terms[edit]
There are also many other descriptive terms, often with very specialised usage and confined to specific taxonomic groups.[37]
- Channelled
Veins sunken below the surface, resulting in a rounded channel
- Guttered
Veins partly prominent, the crest above the leaf lamina surface, but with channels running along each side, like gutters
- Obscure
Veins not visible, or not at all clear; if unspecified, then not visible with naked eye
- Plinervy (plinerved)
More than one main vein (nerve) at the base. Lateral secondary veins branching from a point above the base of the leaf. Usually expressed as a suffix, as in 3-plinerved or triplinerved leaf. In a 3-plinerved (triplinerved) leaf three main veins branch above the base of the lamina (two secondary veins and the main vein) and run essentially parallel subsequently, as in Ceanothus and in Celtis. Similarly a quintuplinerve (five-veined) leaf has four secondary veins and a main vein. A pattern with 3-7 veins is especially conspicuous in Melastomataceae. The term has also been used in Vaccinieae. The term has been used as synonymous with acrodromous, palmate-acrodromous or suprabasal acrodromous, and is thought to be too broadly defined.[38][38]
- Scalariform
Veins arranged like the rungs of a ladder, particularly higher order veins
- Submarginal
veins running close to leaf margin
- Trinerved
2 major basal nerves besides the midrib
The terms megaphyll, macrophyll, mesophyll, notophyll, microphyll, nanophyll and leptophyll are used to describe leaf sizes (in descending order), in a classification devised in 1934 by Christen C. Raunkiær and since modified by others.[39]
Anatomy (medium and small scale)[edit]
Medium-scale features[edit]
Leaves are normally extensively vascularised and typically have networks of vascular bundles containing xylem, which supplies water for photosynthesis, and phloem, which transports the sugars produced by photosynthesis. Many leaves are covered in trichomes (small hairs) which have diverse structures and functions.
Small-scale features[edit]
The major tissue systems present are
- The epidermis, which covers the upper and lower surfaces
- The mesophyll tissue inside the leaf, which is rich in chloroplasts (also called chlorenchyma)
- The arrangement of veins (the vascular tissue)
These three tissue systems typically form a regular organisation at the cellular scale. Specialised cells that differ markedly from surrounding cells, and which often synthesise specialised products such as crystals, are termed idioblasts.[40]
Major leaf tissues[edit]
The epidermis is the outer layer of cells covering the leaf. It is covered with a waxy cuticle which is impermeable to liquid water and water vapor and forms the boundary separating the plant's inner cells from the external world. The cuticle is in some cases thinner on the lower epidermis than on the upper epidermis, and is generally thicker on leaves from dry climates as compared with those from wet climates.[citation needed] The epidermis serves several functions: protection against water loss by way of transpiration, regulation of gas exchange, secretion of metabolic compounds, and (in some species)[which?] 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 tissue includes several differentiated cell types; epidermal cells, epidermal hair cells (trichomes), cells in the stomatal complex; guard cells and subsidiary cells. The epidermal cells are the most numerous, largest, and least specialized and form the majority of the epidermis. These are typically more elongated in the leaves of monocots than in those of dicots.
Chloroplasts are generally absent in epidermal cells, the exception being the guard cells of the stomata. The stomatal pores perforate the epidermis and are surrounded on each side by chloroplast-containing guard cells, and two to four subsidiary cells that lack chloroplasts, forming a specialized cell group known as the stomatal complex. The opening and closing of the stomatal aperture is controlled by the stomatal complex and regulates the exchange of gases and water vapor between the outside air and the interior of the leaf. Stomata therefore play the important role in allowing photosynthesis without letting the leaf dry out. In a typical leaf, the stomata are more numerous over the abaxial (lower) epidermis than the adaxial (upper) epidermis and are more numerous in plants from cooler climates.
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".
In ferns and most flowering plants, the mesophyll is divided into two layers:
- An upper palisade layer of vertically elongated cells, one to two cells thick, directly beneath the adaxial epidermis, with intercellular air spaces between them. Its cells contain many more chloroplasts than the spongy layer. 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.[citation needed] 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.
- Beneath the palisade layer is the spongy layer. The cells of the spongy layer are more branched and not so tightly packed, so that there are large intercellular air spaces between them for oxygen and carbon dioxide to diffuse in and out of during respiration and photosynthesis. These cells contain fewer chloroplasts than those of the palisade layer. The pores or stomata of the epidermis open into substomatal chambers, which are connected to the air spaces between the spongy layer cells.
Leaves are normally green, due to chlorophyll in chloroplasts in the chlorenchyma cells. Plants that lack chlorophyll cannot photosynthesize.
Vascular tissue[edit]
The veins are the vascular tissue of the leaf and are located in the spongy layer of the mesophyll. The pattern of the veins is called venation. In angiosperms the venation is typically parallel in monocotyledons and forms an interconnecting network in broad-leaved plants. They were once thought to be typical examples of pattern formation through ramification, but they may instead exemplify a pattern formed in a stress tensor field.[41][42][43]
A vein is made up of a vascular bundle. At the core of each bundle are clusters of two distinct types of conducting cells:
- Xylem: cells that bring water and minerals from the roots into the leaf.
- Phloem: cells that usually move sap, with dissolved sucrose(glucose to sucrose) produced by photosynthesis in the leaf, out of the leaf.
- A sheath of ground tissue made of lignin surrounding the vascular tissue. This sheath has a mechanical role in strengthening the rigidity of the leaf.
The xylem typically lies on the adaxial side of the vascular bundle and the phloem typically lies on the abaxial side. Both are embedded in a dense parenchyma tissue, called the sheath, which usually includes some structural collenchyma tissue.
Leaf development[edit]
According to Agnes Arber's partial-shoot theory of the leaf, leaves are partial shoots,[44] being derived from leaf primordia of the shoot apex.[8] Compound leaves are closer to shoots than simple leaves. Developmental studies have shown that compound leaves, like shoots, may branch in three dimensions.[45][46] On the basis of molecular genetics, Eckardt and Baum (2010)[citation needed] concluded that "it is now generally accepted that compound leaves express both leaf and shoot properties."
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 boundary layer reducing water loss.
- Waxy leaf surfaces reduce water loss.
- Large surface area provides large area for sunlight and shade for plant to minimize heating and reduce water loss.
- In harmful levels of sunlight, specialised leaves, opaque or partly buried, admit light through a translucent leaf window for photosynthesis at 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 phytoliths in grasses, raphides in Araceae).
- Petals attract pollinators.
- Spines protect the plants (e.g. cacti).
- Special leaves on carnivorous plants are adapted to trapping food, mainly invertebrate prey, though some species trap small vertebrates as well (see carnivorous plants).
- Bulbs store food and water (e.g. onions).
- Tendrils allow the plant to climb (e.g. peas).
- Bracts and pseudanthia (false flowers) replace normal flower structures when the true flowers are greatly reduced (e.g. Spurges).
- Spathe.
Seasonal leaf loss[edit]
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. When the leaf is shed, it leaves a leaf scar 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.[47] Optical masking of chlorophyll by anthocyanins reduces risk of photo-oxidative damage to leaf cells as they senesce, which otherwise may lower the efficiency of nutrient retrieval from senescing autumn leaves.[48]
Interactions with other organisms[edit]
Although not as nutritious as other organs such as fruit, leaves provide a food source for many organisms. The leaf is a vital source of energy production for the plant, and plants have evolved protection against animals that consume leaves, such as tannins, chemicals which hinder the digestion of proteins and have an unpleasant taste. Animals that are specialized to eat leaves are known as folivores.
Some species have cryptic adaptations by which they use leaves in avoiding predators. For example, the caterpillars of some leaf-roller moths will create a small home in the leaf by folding it over themselves. Some sawflies similarly roll the leaves of their food plants into tubes. Females of the Attelabidae, so-called leaf-rolling weevils, lay their eggs into leaves that they then roll up as means of protection. Other herbivores and their predators mimic the appearance of the leaf. Reptiles such as some chameleons, and insects such as some katydids, also mimic the oscillating movements of leaves in the wind, moving from side to side or back and forth while evading a possible threat.
See also[edit]
- Glossary of leaf morphology
- Glossary of plant morphology:Leaves
- Crown (botany)
- Evolution of leaves
- Leaf Area Index
- Leaf protein concentrate
- Leaf sensor – a device that measures the moisture level in plant leaves
- Leaf shape
- Vernation – sprouting of leaves, also the arrangement of leaves in the bud
- ^ a b Esau 2006.
- ^ Cutter 1969.
- ^ Haupt 1953.
- ^ a b Mauseth 2009.
- ^ James et al 1999.
- ^ a b c Stewart & Rothwell 1993.
- ^ Cooney-Sovetts & Sattler 1987.
- ^ a b Simpson 2011, p. 356.
- ^ Krogh 2010.
- ^ Purcell 2016.
- ^ Willert et al 1992.
- ^ Bayer 1982.
- ^ Marloth, Rudolf. "The Flora of South Africa" 1932 Pub. Cape Town: Darter Bros. London: Wheldon & Wesley.
- ^ James, S. A.; Bell, D. T. (2000). "Influence of light availability on leaf structure and growth of two Eucalyptus globulus ssp. globulus provenances". Tree Physiology. 20 (15): 1007–1018. doi:10.1093/treephys/20.15.1007. PMID 11305455.
- ^ Heywood, V.H.; Brummitt, R.K.; Culham, A.; Seberg, O. (2007). Flowering plant families of the world. New York: Firefly books. p. 287. ISBN 9781554072064.
- ^ Hallé, F. (1977). "The longest leaf in palms?". Principes. 21: 18.
- ^ Rolland-Lagan et al 2009.
- ^ a b c Walls 2011.
- ^ a b c Dickison 2000.
- ^ a b Rudall 2007.
- ^ a b c d e f g h Simpson 2011, Leaf venation pp. 465–468
- ^ a b c d Sack & Scoffoni 2013.
- ^ a b c Roth-Nebelsick et al 2001.
- ^ Ueno et al 2006.
- ^ Runions et al 2005.
- ^ Massey & Murphy 1996, Surface-Venation-Texure
- ^ Kew Glossary: Hysteranthous
- ^ Kew Glossary: Synanthous
- ^ Ettingshausen 1861.
- ^ Hickey 1973.
- ^ Hickey & Wolfe 1975.
- ^ Hickey 1979.
- ^ Melville 1976.
- ^ a b Leaf Architecture Working Group 1999.
- ^ Judd et al 2007.
- ^ Florissant Leaf Key 2016.
- ^ Neotropikey 2017.
- ^ a b Pedraza-Peñalosa 2013.
- ^ Whitten, Tony (1996). Ecology of Java and Bali. p. 505. ISBN 9789625930725. Retrieved 18 January 2016.
- ^ Cote, G. G. (2009). "Diversity and distribution of idioblasts producing calcium oxalate crystals in Dieffenbachia seguine (Araceae)". American Journal of Botany. 96 (7): 1245–54. doi:10.3732/ajb.0800276. PMID 21628273.
- ^ Couder, Y.; Pauchard, L.; Allain, C.; Adda-Bedia, M.; Douady, S. (1 July 2002). "The leaf venation as formed in a tensorial field". The European Physical Journal B. 28 (2): 135–138. Bibcode:2002EPJB...28..135C. doi:10.1140/epjb/e2002-00211-1.
- ^ Corson, Francis; Adda-Bedia, Mokhtar; Boudaoud, Arezki (2009). "In silico leaf venation networks: Growth and reorganization driven by mechanical forces". Journal of Theoretical Biology. 259 (3): 440–448. doi:10.1016/j.jtbi.2009.05.002. PMID 19446571.
- ^ Laguna, Maria F.; Bohn, Steffen; Jagla, Eduardo A.; Bourne, Philip E. (2008). "The Role of Elastic Stresses on Leaf Venation Morphogenesis". PLoS Computational Biology. 4 (4): e1000055. Bibcode:2008PLSCB...4E0055L. doi:10.1371/journal.pcbi.1000055. PMC 2275310. PMID 18404203.
- ^ Arber, A. (1950) The Natural Philosophy of Plant Form. Cambridge University Press.
- ^ Rutishauser, R. and Sattler, R. 1997. Expression of shoot processes in leaf development of Polemonium caeruleum. Botanische Jahrbücher für Systematik 119: 563-582.
- ^ Lacroix, C.; Jeune, B.; Purcell-Macdonald, S. (2003). "Shoot and compound leaf comparisons in eudicots: Dynamic morphology as an alternative approach". Botanical Journal of the Linnean Society. 143 (3): 219–230. doi:10.1046/j.1095-8339.2003.00222.x.
- ^ Thomas F. Döring; Marco Archetti; Jim Hardie (2009), "Autumn leaves seen through herbivore eyes", Proceedings of the Royal Society B: Biological Sciences, 276 (1654): 121–127, doi:10.1098/rspb.2008.0858, PMC 2614250, PMID 18782744
- ^ Feild, T. S.; Lee, D. W.; Holbrook, N. M. (2001). "Why leaves turn red in autumn. The role of anthocyanins in senescing leaves of red-osier dogwood". Plant Physiology. 127 (2): 566–74. doi:10.1104/pp.010063. PMC 125091. PMID 11598230.
Books and chapters[edit]
- Bayer, M. B. (1982). The New Haworthia Handbook. Kirstenbosch: National Botanic Gardens of South Africa. ISBN 0-620-05632-0.
- Cutter, E.G. (1969). Plant Anatomy, experiment and interpretation, Part 2 Organs. London: Edward Arnold. p. 117. ISBN 0713123028.
- Dickison, William C. (2000). Integrative Plant Anatomy. Academic Press. ISBN 978-0-08-050891-7.
- Esau, Katherine (2006) [1953]. Evert, Ray F, ed. Esau's Plant Anatomy: Meristems, Cells, and Tissues of the Plant Body: Their Structure, Function, and Development (3rd. ed.). New York: John Wiley & Sons Inc. ISBN 9780470047378.
- Ettingshausen, C. (1861). Die Blatt-Skelete der Dicotyledonen mit besonderer Ruchsicht auf die Untersuchung und Bestimmung der fossilen Pflanzenreste. Vienna: Classification of the Architecture of Dicotyledonous.
- Haupt, Arthur Wing (1953). Plant morphology. McGraw-Hill.
- Hickey, LJ. A revised classification of the architecture of dicotyledonous leaves. pp. i 5–39., in Metcalfe & Chalk (1979)
- Judd, Walter S.; Campbell, Christopher S.; Kellogg, Elizabeth A.; Stevens, Peter F.; Donoghue, Michael J. (2007) [1st ed. 1999, 2nd 2002]. Plant systematics: a phylogenetic approach (3rd ed.). Sinauer Associates. ISBN 0-87893-407-3.
- Krogh, David (2010), Biology: A Guide to the Natural World (5th ed.), Benjamin-Cummings Publishing Company, p. 463, ISBN 978-0-321-61655-5
- Leaf Architecture Working Group (1999). "Manual of Leaf Architecture - morphological description and categorization of dicotyledonous and net-veined monocotyledonous angiosperms" (PDF). Smithsonian Institution. ISBN 0-9677554-0-9.
- Mauseth, James D. (2009). Botany: an introduction to plant biology (4th ed.). Sudbury, Mass.: Jones and Bartlett Publishers. ISBN 978-0-7637-5345-0.
- Metcalfe, CR; Chalk, L, eds. (1979) [1957]. Anatomy of the Dicotyledons: Leaves, stem and wood in relation to taxonomy, with notes on economic uses. 2 vols. (2nd ed.). Oxford: Clarendon Press. ISBN 978-0-19-854383-1.
- Prance, Ghillean Tolmie (1985). Leaves: the formation, characteristics and uses of hundreds of leaves found in all parts of the world. Photographs by Kjell B. Sandved. London: Thames and Hudson. ISBN 0-500-54104-3.
- Rudall, Paula J. (2007). Anatomy of flowering plants: an introduction to structure and development (3rd ed.). Cambridge: Cambridge University Press. ISBN 9780521692458.
- Simpson, Michael G. (2011). Plant Systematics. Academic Press. ISBN 0-08-051404-9.
- Stewart, Wilson N; Rothwell, Gar W. (1993) [1983]. Paleobotany and the Evolution of Plants (2nd ed.). Cambridge University Press. ISBN 978-0-521-38294-6.
- Willert, Dieter J. von; Eller, BM; Werger, MJA; Brinckmann, E; Ihlenfeldt, H-D (1992). Life Strategies of Succulents in Deserts: With Special Reference to the Namib Desert. CUP Archive. ISBN 978-0-521-24468-8.
- Cooney-Sovetts, C.; Sattler, R. (1987). "Phylloclade development in the Asparagaceae: An example of homoeosis". Botanical Journal of the Linnean Society. 94 (3): 327–371. doi:10.1111/j.1095-8339.1986.tb01053.x.
- Hickey, Leo J. (1 January 1973). "Classification of the Architecture of Dicotyledonous Leaves" (PDF). American Journal of Botany. 60 (1): 17–33. doi:10.2307/2441319.
- Hickey, Leo J.; Wolfe, Jack A. (1975). "The Bases of Angiosperm Phylogeny: Vegetative Morphology". Annals of the Missouri Botanical Garden. 62 (3): 538. doi:10.2307/2395267.
- Shelley, A.J.; Smith, W.K.; Vogelmann, T.C. (1998). "Ontogenetic differences in mesophyll structure and chlorophyll distribution in Eucalyptus globulus ssp. globulus (Myrtaceae)". American Journal of Botany. 86 (2): 198–207. doi:10.2307/2656937. PMID 21680359.
- Melville, R. (November 1976). "The Terminology of Leaf Architecture". Taxon. 25 (5/6): 549. doi:10.2307/1220108.
- Pedraza-Peñalosa, Paola; Salinas, Nelson R.; Wheeler, Ward C. (26 April 2013). "Venation patterns of neotropical blueberries (Vaccinieae: Ericaceae) and their phylogenetic utility" (PDF). Phytotaxa. 96 (1): 1. doi:10.11646/phytotaxa.96.1.1.
- Rolland-Lagan, Anne-Gaëlle; Amin, Mira; Pakulska, Malgosia (January 2009). "Quantifying leaf venation patterns: two-dimensional maps". The Plant Journal. 57 (1): 195–205. doi:10.1111/j.1365-313X.2008.03678.x.
- Roth-Nebelsick, A; Uhl, Dieter; Mosbrugger, Volker; Kerp, Hans (May 2001). "Evolution and Function of Leaf Venation Architecture: A Review". Annals of Botany. 87 (5): 553–566. doi:10.1006/anbo.2001.1391.
- Runions, Adam; Fuhrer, Martin; Lane, Brendan; Federl, Pavol; Rolland-Lagan, Anne-Gaëlle; Prusinkiewicz, Przemyslaw (1 January 2005). "Modeling and Visualization of Leaf Venation Patterns". ACM SIGGRAPH 2005 Papers. ACM: 702–711. doi:10.1145/1186822.1073251.
- Sack, Lawren; Scoffoni, Christine (June 2013). "Leaf venation: structure, function, development, evolution, ecology and applications in the past, present and future". New Phytologist. 198 (4): 983–1000. doi:10.1111/nph.12253.
- Ueno, Osamu; Kawano, Yukiko; Wakayama, Masataka; Takeda, Tomoshiro (1 April 2006). "Leaf Vascular Systems in C3 and C4 Grasses: A Two-dimensional Analysis". Annals of Botany. 97 (4): 611–621. doi:10.1093/aob/mcl010.
- Walls, R. L. (25 January 2011). "Angiosperm leaf vein patterns are linked to leaf functions in a global-scale data set". American Journal of Botany. 98 (2): 244–253. doi:10.3732/ajb.1000154.
- "Leaves"., in Massey & Murphy (1996)
- Massey, Jimmy R.; Murphy, James C. (1996). "Vascular plant systematics". NC Botnet. University of North Carolina at Chapel Hill. Retrieved 19 January 2016.
- Geneve, Robert. "Leaf" (PDF). PLS 220: Intoduction to plant identification. University of Kentucky: Department of Horticulture.
- "Florissant Fossil Beds Leaf Key" (PDF). Florissant Fossil Beds National Monument. National Parks Service, US Department of the Interior. Retrieved 16 February 2017.
- Purcell, Adam (16 January 2016). "Leaves". Basic Biology. Adam Purcell. Retrieved 17 February 2017.
- "Glossary of botanical terms". Neotropikey. Royal Botanic Gardens, Kew. Retrieved 18 February 2017.
External links[edit]
- Ernest Ingersoll (1920). "Leaves". Encyclopedia Americana.
- Science aid: Leaf Leaf structure and transpiration resource for teens.
- Cleared Leaves DB An open database for cleared leaves with full annotation.
|Wikimedia Commons has media related to Leaves.
|Look up leaf in Wiktionary, the free dictionary.


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Hotfrog US provides information regarding Roy D Steinberg in Voorhees NJ. Roy D Steinberg is located at 26 Old Stevens Ln and provides Healthcare,Mental Health,Psychologist services. Contact them on (856) 782-1943 or by …
Union, NJ
Our highly trained and sensitive caregivers combine a special blend of professional expertise with compassionate attention to the unique needs of senior adults and their families
elderly home health careSenior Care ManagementRespite Carehome care services
Mount Laurel, NJ
A+ Senior Care provides specialized quality in-home care to seniors. We are passionate about senior care. Caregivers are insured, bonded and have been criminally checked. An A+ Senior Care representative is available an…
Medication RemindersNon-medical Home CareSenior CaregiverSenior Home Care
Freehold, NJ
Quality Assisted Living, Caregivers, Senior Care and In Home Care from Affordable Senior Home Care. Serving Toms River, NJ and Ocean County area. 1-866-377-2903
Home Health Care CompanySenior Home CareIn Home Care
New Brunswick, NJ
The Care Street mission is straightforward and compelling: to provide a comprehensive range of the highest quality home health services in a professional and efficient manner that will enhance our clients' quality of lif…
Bordentown, NJ
Our system empowers you to select your caregiver – you will have the opportunity to interview the caregivers we recommend Our caring and affectionate caregivers provide up to 24 hour care in the comfort and fam…
Elderly CareSenior Care
Bernardsville, NJ
We offer an array of personalized services from in-home (included assisted living and other facilities) and live-in care for the older adult to "doula" support for the new mother. We pride ourselves on having the best an…
Nutley, NJ
We are known for making a difference in the lives of seniors that are in our care while remaining in the comfort of their own home. We treat and care for our clients like they were part of our own family. Our services in…
Non-medical Home CareSenior Home CareIn Home CareEldercare
Morris Plains, NJ
Family First HomeCare - We care for NJ Seniors at Home. Certified Caregivers to care for the elderly Family First HomeCare is the perfect solution for seniors and others who aren't ready to leave their homes for an i…
Family CaregiverCare For The ElderlyHomecareSenior Care
Select Care is licensed by the Division of Consumer Affairs, and all of our staff are certified, insured, and undergo an intensive interview, health screening, and background checks We are licensed and insured …
Personal Care ServicesCourteous Staff Members
Piscataway, NJ
Caring Matters provides in-home high quality, compassionate care you can trust to adults; and seniors who need non-medical care assistance with daily activities. Our home care services consisting of Licensed Registered …
Aging CarePersonal Care Assistanthome health aidesRespite Services
Tenafly, NJ
The ComedyCures Foundation tickles funny bones Through large & small scale therapeutic comedy programs, we entertain and educate patients, families and caregivers about the power of a comic perspective and the p…
Therapeutic Humor
Morristown, NJ
Bridged Care bridges the gap between health care professionals, as the point of care, with loved ones and family members. For Caregivers, our online and mobile interface makes it possible to keep the families of your cli…
Through large & small scale therapeutic comedy programs, we entertain and educate patients, families and caregivers about the power of a comic perspective and the positive benefits of laughter on the body, mind, and…
Fox GERI has received 5013 status from the IRS All monetary contributions to Fox GERI are now tax deductible Our mission is to provide high quality educational services to the public on health Please …
Assisted LivingOccupational And Speech Therapists
Hasbrouck Heights, NJ
Phillipsburg, NJ
New caregiver candidates are rigorously interviewed in person to determine their eligibility as a Crossroads Homecare caregiver We believe the right combination of experience, skills and attitude can overcome e…
Household ChoresHouseworkElder CareHome Management
Whippany, NJ
Seniors and Caregivers also love our where they can find out about local events and activities, many of which are free, all of which are worthwhile The Hersch family truly appreciates and values the opportunity…
Union County, NJ
We only employ the best caregivers available We strive for your peace of mind and satisfaction by selecting the caregiver, with the proper skills and personality to fit our client's needs
Medical Home CareHomecare ServicesIn Home CareHomecare
ComedyCures was honored for our outstanding impact on patients' lives Payaso Entertainment and many talented comedians joined forces on 6 television comedy specials to further support ComedyCures programs …
Inspire connects patients, families, friends, caregivers and health professionals for health and wellness support. Connect with others who know what you're going through
Manalapan, NJ
Our caregivers will work with your loved one on exercise routines to maintain flexibility and dexterity, monitor nutritional intake to maintain strength, help them with daily activities of living, and adhere to the …
Senior Home CareIn Home CareElder CareAlzheimer's care
Old Bridge, NJ
Health Senior ServicesSenior Center TransportationAssisted Living HomecarePediatric Home Healthcare
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Is Bank Muamalat @ Klang in No. 46 & 48, Jalan Kelicap 42A/Ku1, Klang Bandar Diraja, Off Jalan Meru, 41050, Klang, Selangor, Klang Selangor 41150 your business? Claim your listing and attract more leads by adding more content, photos and other business details.
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Hampton Inn & Suites Craig
Get an overview of this hotelHotel in Craig with free breakfast and indoor pool
Situated in Craig, this hotel is within 3 mi (5 km) of Craig City Hall, Sandrocks Nature Trail and Petroglyphs, and Museum of Northwest Colorado. Marcia Car and Craig City Park are also within 3 mi (5 km).
Hotel Features
An indoor pool, a fitness center, and a spa tub are available at this smoke-free hotel. Free buffet breakfast, free WiFi in public areas, and free self parking are also provided. Additionally, a business center, meeting rooms, and a 24-hour front desk are onsite.
Room Amenities
All 89 rooms offer free WiFi, TVs, and hair dryers. Ironing boards and desks are also available to guests.Information missing or incorrect? Tell us!Opens in a new window
Hampton Inn & Suites Craig
Hotel Amenities
Hotel Amenities
In addition to an indoor pool, Hampton Inn & Suites Craig provides a spa tub and a fitness center. Public areas are equipped with complimentary high-speed wireless Internet access. Business-related amenities consist of a business center and meeting rooms. Guests can enjoy a complimentary breakfast. 4 meeting rooms are available. This Craig hotel also offers express check-out and a safe deposit box at the front desk. Complimentary self parking is available on site.
Hampton Inn & Suites Craig is a smoke-free property.
- 24-hour front desk
- Total number of rooms - 89
- Number of floors - 4
- Free breakfast
- Business center
- Express check-out
- Fitness facilities
- Free WiFi
- Meeting rooms
- Safe-deposit box at front desk
- Number of meeting rooms - 4
- Free self parking
- Smoke-free property
- Number of buildings/towers - 1
- Spa tub
- Indoor pool
Family Friendly Amenities
- Family Hotel
- Indoor pool
- Free Wi-Fi
- Rollaway/extra beds available
- Refrigerator (on request)
- Microwave (on request)
- Cribs/infant beds (complimentary)
Available in all rooms: Free WiFi
Available in some public areas: Free WiFi
Free self parking
Room Amenities
- Microwave (on request)
- Refrigerator (on request)
- Private bathroom
- Hair dryer
- Bathtub or shower
- Iron/ironing board
- Television
- Desk
- Rollaway/extra beds available
- Free WiFi
- Connecting/adjoining rooms available
- Free cribs/infant beds
Where to Eat
Guests are offered a complimentary hot/cold buffet breakfast.
Nearby Things to Do
An indoor pool and a spa tub are on site. Other recreational amenities include a fitness center.
If you have requests for specific accessibility needs, please note them at check-out when you book your room.
- Accessible bathroom
- In-room accessibility
Hotel Policies
Check-in time starts at 4 PM
Check-out time is 11 AM
Payment types
Children and extra beds
- Children are welcome.
- Rollaway/extra beds are available.
- Free cribs (infant beds)!
- Up to 60 lb
You need to know
Extra-person charges may apply and vary depending on property policy.
Government-issued photo identification and a credit card or cash deposit are required at check-in for incidental charges.
Special requests are subject to availability upon check-in and may incur additional charges. Special requests cannot be guaranteed.
Hotel Name
- Hampton Inn Craig
- Hampton Inn Hotel Craig
- Hampton Inn & Suites Craig Hotel Craig
- Hampton Inn And Suites Craig
We should mention
The property has connecting/adjoining rooms, which are subject to availability and can be requested by contacting the property using the number on the booking confirmation.


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Yo Gabba Gabba! Tour Sells Out In New York
By Ray Waddell
Yo Gabba Gabba! Live: There's a Party in My City, the first live U.S. touring shows for the popular children's program "Yo Gabba Gabba!," looks like a hit.
Conceived by animation/entertainment company W!ldbrain and produced and promoted by Michael Cohl's Miami-based S2BN Entertainment, YGG Live put six shows in three cities -- New York, Chicago and Los Angeles -- on sale today (Oct. 9) and immediately sold out New York's two shows and is considering adding more. Chicago and Los Angeles go up today as well.
The tour begins Nov. 14 in Los Angeles at the Shrine Expo Hall. New York's Beacon Theatre is Nov. 21, and the Rosemont Theatre outside of Chicago is Dec. 5. Former artist manager Mike Luba, president of music and family entertainment at S2BN, is running point for the company on the Yo Gabba Gabba! tour.
The New York blowout for a tour untested in North America is an impressive debut. "We didn't really know, this being the first time out," Luba admits. "We weren't sure if it was something we should book into Brooklyn Bowl and see if 800 people show up, or should we go into the Garden and see if 10,000 show up. We feel like we kind of settled into the right spot, and we're going to give people a really good experience."
YGG as a family show tour jumps into a lucrative but competitive space that annually fields such long-time winners as Dora the Explorer, Sesame Street Live and Clifford the Big Red Dog, among others. Luba believes the rock sensibilities of YGG that appeals to parents, along with the expertise at S2BN, come at this entertainment niche from another angle.
"Nothing against any of those shows, they all do what they do really well," Luba says of the competition in the family show arena. "We'll take the skills we've learned in doing everything from Broadway to rock concerts to events and exhibitions and put it to use here. It's up to us to not only let people know the show's coming, but to educate [consumers] as to what it's all about so they know what to expect, and to blow their minds when they get there."
S2BN Entertainment bills itself as "a diversified, international family show and event entertainment company specializing in the acquisition, development and production of high caliber touring exhibitions, unique live music tours and events, theatrical performances, and consumer and multimedia product offerings." Cohl is best known as the promoter of the Rolling Stones' world tours since 1989, and was a key player in orchestrating Live Nation's megadeals with such artists as Madonna, U2, Jay-Z and Shakira. Cohl resigned as chairman of the board of Live Nation and vacated his post as CEO of Live Nation Artists in June of 2008, and launched S2BN later that year.
S2BN projects to date include the Bodies Exhibition, FuerzaBruta and the Pete Seeger Clearwater Concert: Creating the Next Generation of Environmental Leaders in celebration of his 90th Birthday. Now it's Yo Gabba Gabba! "It seems like the people out there really want it," Luba says. "I was sitting with [Cohl] one day and said, 'I'm certainly prone to being wrong but I would have to be as wrong about this as anything I've ever done in my life.'"
The show will feature DJ Lance Rock and the YGG characters Brobee, Foofa, Muno, Plex, and Toodee, as well as surprise guests. Luba says the show features very high production values in light, sound and video. The tour comes on the heels of the first YGG album release, "Yo Gabba Gabba! Music Is Awesome," set for release Oct. 20 on Filter Recordings.


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We are women physicians caring for women patients in all phases of life. From the moment that you walk into our tranquil, comfortable reception area, you'll know that you are in a different kind of Ob-Gyn practice-one where you are greeted with warmth and compassion by our staff. We strive to make a personal connection with each patient, to put you at ease, to help you to feel that you are not just in for another "gyno exam". Our providers are committed to getting to know you as an individual through careful listening so that we can work together towards your best health ever. When medical issues arise, we blend the latest medical science, patient education and your unique needs to create treatment options with you. We offer a wide range of services from maternity care and delivery to in-office and hospital surgery. Our practice provides on-site ultrasound and is one of few AIUM-certified facilities in the Denver metro area. Our office is located just south of the DTC, close to the Dry Creek light rail station. We offer appointment times that are convenient for today's busy women. We are passionate about your comfort, tranquility and safety. We feel privileged to be part of each and every one of your lives, to be able to share your greatest joys and to receive your trust. We know that women are the heart of the future.Read more
For the best healthcare for your needs, choose a doctor who specializes in your medical condition.
Board certification indicates that a doctor is highly qualified in the medical field in which they practice.
Check to see that this provider treats your medical condition.
Check to see that this provider performs the procedure that you need.
Check to see if your provider has any malpractices, board actions, or sanctions.
Learn more about where this provider went to medical school, residency, and more.
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Likelihood of recommending Dr. Russell to family and friends is 4.1 out of 5
Obstetrics & Gynecology Specialist


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We are 107,000+ car owners sharing real-world car information.Join Us
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1 comment
|A Clownshoe with the Best Hips in the Business
2002 BMW Z3 Review
The M coupe is a little known number that BMW had no plans to produce until after a group of skunkworks engineers, craving a stiffer chassis, slapped a roof on an M roadster, and barely convinced them was good for the company's performance image. As such, while the basic design,...
|2000 BMW Z3 Review
A member in Alabama, United States
Leg/head room was lacking. Interior seemed cramped for my size.
|2001 BMW Z3 Review
|2000 BMW Z3 Review
A member in Texas, United States
Styling is a personal issue and subjective, but I always liked the looks of the Z3, especially the convertible.
|2002 BMW Z3 Review
Perfection. It has been my dream car since it first came out in 1996.
|2000 BMW Z3 Review
Slick, wide hips and sexy
|2000 BMW Z3 Review
Car of the year in 2000. Best Consumer Reports value for 5 years in a row for its class. � Model Rating - Overall 4 stars � Trim Rating - Overall 5 stars � Appearance 5 stars � Comfort 5 stars � Performance 5 stars � Reliability 5 stars
|2002 BMW Z3 Review
At least 7000$ more than a similarly equipped Miata, for no gain.
Thanks for your interest in TrueDelta!


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Lobster Rumaki
23 Jan 2009
|Cooked lobster
|1 Pound
|Water chestnuts
|1 Cup (16 tbs)
|Soy sauce
|1 Tablespoon
|Bacon slices
|1 Cup (16 tbs)
"Split center of each piece of lobster; insert thin slice of water chestnut.
Brush well with oil and soy sauce.
Wrap in bacon; secure with wooden pick.
Broil until bacon is crisp."
Recipe Summary
Difficulty Level:Easy
Course:Side Dish
Rate It
Nutrition Rank
Nutrition Facts
Serving size: Complete recipe
Calories 1053 Calories from Fat 333
% Daily Value*
Total Fat 38 g59.1%
Saturated Fat 12 g59.8%
Trans Fat 0 g
Cholesterol 471.8 mg157.3%
Sodium 3089.4 mg128.7%
Total Carbohydrates 26 g8.6%
Dietary Fiber 1.3 g5.1%
Sugars 0.3 g
Protein 139 g277.6%
Vitamin A 1.8% Vitamin C 17.8%
Calcium 28.9% Iron 43.6%
*Based on a 2000 Calorie diet
Lobster Rumaki Recipe


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Created attachment 580057 [details]
Uncloseable inspector on top of google.ca
User Agent: Mozilla/5.0 (Macintosh; Intel Mac OS X 10_6_7) AppleWebKit/535.2 (KHTML, like Gecko) Chrome/15.0.874.121 Safari/535.2
Steps to reproduce:
Aurora version 10.0a2
1. Loaded a page
2. Right click -> Inspect Element
3. Right click -> Inspect Element again
4. Exit the inspector using X on the inspector bar.
Actual results:
After step 3, a new inspector appears on top of the old one.
After step 4, the inspector bar disappears but the screen remains greyed out and the inspector box and bubble remains visible. Reloading the page doesn't help. Running Inspector again doesn't help. Visiting another page causes the screen to remain greyed out with the inspector box and bubble.
Expected results:
After step 3, I expect the element to appear in the existing inspector.
After step 4, I expect the inspector to disappear.
I've only been able to reproduce this problem using a confidential internal application. I can visit another page after step 4, and I will attach a screenshot of what that looks like (and another with 2 inspectors.)
Created attachment 580058 [details]
Two inspectors on top of google.ca
Can you also test the latest nightly? http://nightly.mozilla.org/
(In reply to Tim (fmdeveloper) from comment #2)
> Can you also test the latest nightly? http://nightly.mozilla.org/
Unfortunately I can no longer reproduce this in Aurora (which updated itself) or in the latest nightly. It was difficult to reproduce to begin with so I'm not sure if it's been fixed or if it's just not happening anymore. Feel free to resolve this as WORKSFORME...
Will close - if you are able to reproduce again please do post back!
I am experiencing this exact problem on an internal page that I am working on. I am using the 10.0 release for Windows.
Here is a little more information and a test page where I can reproduce this every time.
Test Page: http://webpages.charter.net/jhargett/firefox_insp_test.html
1. Right click on the INPUT element with the label "Name".
2. Click "Inspect Element" in the context menu.
3. While the Inspector is still open and has the screen "dimmed", right click on the SELECT element with the label "Select".
4. Click the "Inspect Element" option in the context menu.
Instead of moving the bubble to the SELECT element, it opens a new Inspector window on top of the old one (this is noticeable because the screen gets even dimmer.) It can't find the SELECT element so it goes in to Inspector mode where it puts a bubble over whatever element you mouse over. If you close that inspector, the original one stays open with no way to close it. The only solution at this point is to close the tab and start over.
Paul, I can reproduce this on nightly, can you take a look please?
I must be missing something. I can't reproduce. I can't right click on the SELECT element (there's no contextual menu for this element).
Maybe it is an issue with a different OS? I am on WinXP PRO SP3. I am using FF 10.0. Here is a screencast of it happening on my computer:
I can't trigger this on win7(aurora), but can trigger on osx 10 (nightly)
Apparently, Firebug needs to be installed to reproduce this bug.
jonmhargett, do you have Firebug installed?
Yes. I will retest with it disabled.
That was it. I disabled firebug and now the context menu is not available when I right click on a SELECT element. This problem exists with Firebug 1.9.1. So, what is the solution: should Firebug fix this or should you guys make it so that SELECT elements are clickable and the inspector switches to that element?
Looks fixed in Firebug 1.10.
Created attachment 594706 [details] [diff] [review]
patch v1
It's not "fixed" in Firebug 1.10, it's just that the right-click-on-form feature is broken in 1.10.
The problem is that Firebug allows right-click on forms, and this behavior is not supported by the builin inspector yet: we get a call to openInspectorUI(this.target) where target is null. We don't correctly test this
This patch fix this test. I opened bug 724546 to support right click on forms.
This should be landed in Beta and Aurora too.
*** Bug 728491 has been marked as a duplicate of this bug. ***
For what it's worth, this is also an issue on Fedora 14, and I assume, other Linux distributions as well. I would have had a hard time figuring out that it was related to having Firebug installed, so good work, guys.


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Boogaard glad to be back on court
KU women's basketball player Krysten Boogaard played for the first time all year in Wednesday's win over Western Illinois.
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Articles tagged christopher reich
26 Jul 2010 // 7:00 AM
A surgeon doing charity work to make up for the sins of his violent past changes his priorities after learning that his new wife married him for an unfathomable reason.READ more
//Mixed media
Animal Collective "Kinda Bonkers" (Singles Going Steady) // Music
Hayley Kiyoko "Palace" (Justin Caruso Remix) (premiere) // Music
Marty Stuart "Way Out West" (Singles Going Steady) // Music
Katie Herzig "I Want to Make You Proud" (video) (premiere) // Music
//Pm Picks
'Who' Will Be the Next Doctor?
// Channel Surfing
"What shall it be? A Doctor with whip-smart delivery of his lines? A woman who will bewitch before she kicks a Dalek's ass? Oh, the possibilities...READ the article


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This week on The CultCast: You'll laugh, you'll cry — wait till you hear the latest lawsuit aimed right at Apple's biggest products. Plus: Leaked photos of EarPods with a Lightning connector look better than expected; why we may never get third-party Apple Watch faces; Netflix is adding a feature it swore would never happen; and a man builds a vibrant photography career in the world of sports with nothing but his iPhone. Oh, and Val Kilmer dances with a carrot in his derriere. Just hit play already.
Our thanks to Freshbooks for supporting this episode. FreshBooks is the easy-to-use invoicing software designed to help small business owners get organized, save time invoicing and get paid faster. Get started now with a 30-day free trial.


text: Feds To Investigate GM Over Recall, Congress To investigate FedsMatt Hardigree3/12/14 9:30amFiled to: The Morning ShiftTMSRecallsChevy CobaltNHTSA60EditPromoteShare to KinjaToggle Conversation toolsGo to permalink This is The Morning Shift, our one-stop daily roundup of all the auto news that's actually important — all in one place at 9:30 AM. Or, you could spend all day waiting for other sites to parse it out to you one story at a time. Isn't your time more important? Advertisement 1st Gear: Feds Look At GM's Recall Yeah, the GM recall ain't going away, and, as we reported yesterday, now the feds are getting involved. Specifically, the Justice Dept. is going to open an investigation into how GM handled the recall. Advertisement Per The Detroit News:Two people briefed on the matter say that a deputy chief of the office's criminal division reached out to lawyers last week gathering information for a possible subpoena to GM; that is a sign the office is mounting a preliminary investigation. The investigation is focused on whether GM notified NHTSA in a timely fashion about the switch problems.So that's happening and I assume it's all going to end right th– oh wait, what? Advertisement Sponsored 2nd Gear: House To Investigate Feds If you missed it yesterday Congress is ready to jump up and down on GM for how they performed. Also, they'll be looking at NHTSA. Per the Freep: Advertisement U.S. Rep. Fred Upton, R-Michigan, chairman of the House Energy and Commerce Committee, said the committee would examine why GM and the National Highway Traffic Safety Administration failed to act quickly when customers started reporting problems nearly a decade ago.3rd Gear: And In Other News... So Fiat CEO Sergio Marchionne is gonna have to make do with just $5 million, a slight decrease from the year before. Advertisement Advertisement Why? He didn't quite meet certain performance metrics for Fiat. Don't feel bad, he also got the right to receive about $25 million in Fiat shares but hasn't taken them yet.He also didn't take any money for any of his work for Chrysler, although I bet they pay for his Blackberry.4th Gear: Who Would Buy Cars.com? We told you yesterday that Cars.com could be worth $3 billion, but who would buy it? Bloomberg reports that Cox Enterprises/Apax Partners — who already own AutoTrader — are interested in bidding for the site. Advertisement Advertisement Cars.com is currently owned by a company made up of some of America's largest newspaper pubs who, I'm guessing, wouldn't mind the extra income from the sale.5th Gear: Skoda Has A Great Month Skoda, America's favorite Czech automaker, is well on their way to another strong year in an otherwise still weak European market with sales up 4.6% in February to 67,100 vehicles. While their best-selling Skoda Octavia was off a little for the month, and the Suberb was somehow down 23.3% despite being awesome, their Skoda Rapid was up a whopping 212.1% for the month. Advertisement Keep it up Skoda, we love you.Reverse: The Italian Henry Ford Advertisement On this day in 1921, Giovanni "Gianni" Agnelli, the glamorous, powerful Italian business tycoon who turned Fiat, his family's car company, into an international conglomerate, is born in Turin, Italy. Agnelli was named for his grandfather, who founded Fabbrica Italiana Automobili Torino, later known as Fiat, in 1899.[HISTORY] Advertisement Neutral: How badly does this go for GM?Normal bad, Toyota bad? How bad?Photo Credit: AP ImagesReply60 repliesLeave a reply


text:
Call for Assistance Devices
1 result found that includes 3 products
- Call for Assistance Kit Call for Assistance Kit
Multiple product options available
These devices emit an alarm/flashing light to indicate the need for help during a medical emergency. Use in health care facilities, restrooms, and handicap areas.
Multiple product options available
Price $147.80–$189.00
Call for assistance devices from Grainger are designed for areas that require a call-for-help or an emergency switch. Such areas may include health care facilities, public restroom areas and handicapped areas. They include pull cord switches that provide emergency call activation. The call may be reset at the initiating switch. Cords can be replaced with a standard nylon cord.


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I came to new jersey in my quest for a better life. Im getting very comfortable and just lacking a family. Could yours be the one ive dreamed about?
* adoption fee $100
How to Adopt Me!
If you are interested in learning more about me, and possibly making me part of your family, please go to wagoninn.rescuegroups.org. Click on the orange Online Adoption Application button to fill out an application. A Wag On Inn volunteer will be in touch soon!
Thank You!
Your message has been sent to Wag On Inn Rescue.
You'll receive a copy, too, at to help you keep track of which pets you've inquired about, and which shelters and rescues you've emailed.
NOTE: Some shelters have physical locations you can visit; some of these shelters may only have pets for a limited time, so please do not wait for a reply—just go visit the shelter! Other organizations are rescue groups run by busy volunteers who may take a while to reply. You can find information about the shelter or rescue group caring for this pet, and their adoption procedures, on the pet's details page on Adopt-a-Pet.com.


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The dev blog also deals with Albion Online flagging. What is really interesting about this system is that it is not a post-attack penalty like those found in most mmo games. In certain zones, a player will have to flag themselves as a "villain/criminal" before they can attack another player. The difference found here is that the process of flagging can take up to a minute and the player cannot move during that time. This gives other players the chance to flee the area rather than just be ambushed by someone much more powerful. The thought behind this flagging is to cut down on abuse.
You can read the full blog post on Albion Online Power Leveling and the different types of zones here.Albion Online dubs itself as the first, true cross-platform medieval sandbox game. Featuring stunning graphics and a fight for your life, the game is placed in the medieval era with you traversing the dangerous worlds. Along the way, you'll be collecting goods, crafting weapons and armor, hunting, and tending your own farm in an attempt to survive.
An amazing game with a lot of potential, Albion Online won't be out until early 2016, but you can sign up for beta and test the game out for yourself.
Farmers that want to diversify the way that they make money off of the digital world will find that Albion is the perfect candidate for making a little extra money off of your farming. Also, the game has stunning graphics and a lot of fun ways to customize your character. And, with a player-driven economy, you'll have plenty of ways to monetize this game.


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Radiopaq Jazz Review - Page 9
Head-Fi's Best Sellers
My replacement headphones turned up today...
... they sent me the iPhone ones again!*@?
I do not recommend ordering from scienceandtechnology on ebay, I suspect that they do not actually have the "proper" jazz tuned headphone at all.
Edit: DO NOT order from this seller unless you want the iphone version, they do not have any of the normal jazz headphones.
Edited by clothears202 - 3/2/12 at 9:13am
- 87 Posts. Joined 1/2005
- Select All Posts By This User
Those of you who have got the good old version of this (in that Apple-style plastic box) recently, did you buy from seller Allcam/eHome, or directly from Amazon?
Allcam has all versions at £9.98. Amazon has the Classic at £10.06 and the Jazz at £34.20, no idea if Amazon themselves had the Jazz down to £10.
- 525 Posts. Joined 12/2011
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I got mines (the good ones) from amazon.uk at £10.
- 52 Posts. Joined 7/2011
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- 235 Posts. Joined 4/2005
- Location: Greece
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Those of you who have got the good old version of this (in that Apple-style plastic box) recently, did you buy from seller Allcam/eHome, or directly from Amazon?
Allcam has all versions at £9.98. Amazon has the Classic at £10.06 and the Jazz at £34.20, no idea if Amazon themselves had the Jazz down to £10.
I got mine from Allcam/eHome bought through Amazon. They are the good old version!
After much to-ing and fro-ing scienceandtechnology finally admitted that they do not have any of the normal Jazz headphones left. If you order from them you will get a set of the iphone ones (new style with braided cables), although their listing still shows the non-iphone version.
I got a refund and a set of the iphone model by way of an apology.
Just as I was ordering a set of the original version from Amazon I decided to give the ones I had a go. I can't see anything wrong with the way they sound!
Having run an MP3 player through them for 12 hours or so I find them quite satisfactory. I can make out everything that is going on in the music I have tried (including the hiss on some less than perfectly digitised Pink Floyd) without any apparent distortion or blurring of the various instruments. They produce a similar volume level from portable MP3 players to the original earbuds.
They certainly look like alloy to me. They are obviously made in two parts, but there is no step between the black body and the silver front part.
The cable had set into kinks from the way it had been packaged, but I have been able to straighten most of these out fairly easily.
I'm hoping that the slider and shirt clip I have fitted will help sort out the microphonics from the cable.
Most interestingly I discovered that the microphone and button on these iphone headphones work perfectly well on my Android phone (Motorola Atrix running Gingerbread). I can use the button to bring up voice commands or to pause/play MP3s (Winamp), including when it is locked. Very useful since commuting is the main thing I wanted headphones for.
Thanks for putting me onto these headphones guys, hopefully some of the above will help anyone else looking for a set.
Edited by clothears202 - 3/3/12 at 8:20am
I just got these a few days ago and got the new version, i have the old version to and they both sound the same to me...the bass sucked at the start butyou gotta burn them in and the bass greatly improves and the highs tame down a bit.
there 8£ on ebay at sec
EDIT swapped back to old ones these a clear difference in bass, iphones one suck quite a bit i wonder if its the fact the cable is same width and has to carry the mic and buitton wire.
Edited by daniel r - 1/28/13 at 11:00am
- 10,241 Posts. Joined 10/2008
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thats frankly an insane price for them
Yer i got the origs from amazon and there amazing, never seen heard such quality basss with zero distortion.
Edited by daniel r - 1/31/13 at 4:08pm


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Credit Spread Interdependencies of European States and Banks During the Financial Crisis
Adrian Alter
International Monetary Fund
Yves Stephan Schüler
European Central Bank (ECB)
December 10, 2011
We investigate the interdependence of the default risk of several Eurozone countries (France, Germany, Italy, Ireland, Netherlands, Portugal, and Spain) and their domestic banks during the period June 2007 - May 2010, using daily credit default swaps (CDS). Bank bailout programs changed the composition of both banks' and sovereign balance sheets and, moreover, affected the linkage between the default risk of governments and their local banks. Our main findings suggest that in the period before bank bailouts the contagion disperses from bank credit spreads into the sovereign CDS market. After bailouts, a financial sector shock affects more strongly sovereign CDS spreads in the short-run, however, the impact becomes insignificant at a long horizon. Furthermore, government CDS spreads become an important determinant of banks' CDS series. The interdependence of government and bank credit risk is heterogeneous across countries, but homogeneous within the same country.
Number of Pages in PDF File: 38
Keywords: credit default swaps, financial crisis, private-to-public risk transfer, bailout programs, government interventions, cointegration, generalized impulse responses
JEL Classification: C58, G01, G18, G21
Date posted: May 17, 2011 ; Last revised: January 16, 2012
Suggested Citation