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Plant cuticle
From Wikipedia, the free encyclopedia Jump to: navigation, search

Water beads on the waxy cuticle of kale leaves

Anatomy of a leaf

Plant cuticles are a protective waxy covering produced only by the epidermal cells [1] of leaves, young shoots and all other aerial plant organs without periderm. The cuticle tends to be thicker on the top of the leaf, but is not always thicker in xerophytic plants living in dry climates than in mesophytic plants from wetter climates, despite a persistent myth to that effect. The cuticle is composed of an insoluble cuticular membrane impregnated by and covered with soluble waxes. Cutin, a polyester polymer composed of inter-esterified omega hydroxy acids which are cross-linked by ester and epoxide bonds, is the best-known structural component of the cuticular membrane.[2][3] The cuticle can also contain a non-saponifiable hydrocarbon polymer known as Cutan.[4] The cuticular membrane is impregnated with cuticular waxes[5] and covered with epicuticular waxes, which are mixtures of hydrophobic aliphatic compounds, hydrocarbons with chain lengths typically in the range C16 to C36.[6] The plant cuticle is one of a series of innovations, together with stomata, xylem and phloem and intercellular spaces in stem and later leaf mesophyll tissue, that plants evolved more than 450 million years ago during the transition between life in water and life on land.[7] Together, these features enabled plant shoots exploring aerial environments to conserve water by internalising the gas exchange surfaces, enclosing them in a waterproof membrane and providing a variable-aperture control mechanism, the stomatal guard cells, which regulate the rates of transpiration and CO2 exchange. In addition to its function as a permeability barrier for water and other molecules, the micro and nano-structure of the cuticle confer specialised surface properties that prevent contamination of plant tissues with external water, dirt and microorganisms. Many plants, such as the leaves of the sacred lotus (Nelumbo nucifera) exhibit ultra-hydrophobic and selfcleaning properties that have been described by Barthlott and Neinhuis (1997).[8] The lotus effect has potential uses in biomimetic technical materials. "The waxy sheet of cuticle also functions in defense, forming a physical barrier that resists penetration by virus particles, bacterial cells, and the spores or growing filaments of fungi".
[9]

Plant Adaptations to Aquatic Life Totally Submerged Plants

Water Starwort in a marsh pool.

Totally submerged plants are the true water plants or hydrophytes. Because they are truly aquatic they have the greatest number of adaptations to life in water. These include: The presence of little or no mechanical strengthening tissue in stems and leaf petioles. If these plants are removed from the water, they hang limply. They are normally supported by water all around them and so have no need of mechanical strengthening. Indeed, this would be a distinct disadvantage as it would limit flexibility in the event of changes in water level or water movements. Submerged plants lack the external protective tissues required by land plants to limit water loss. The epidermal (outermost) layer shows very little, if any, sign of cuticle formation. All the surface cells appear to be able to absorb water, nutrients and dissolved gases directly from the surrounding water. As a result, the internal system of tubes (xylem) which normally transports water from the roots to all parts of the plant is often greatly reduced, if not absent. Thus, if these plants are removed from the water, they wilt very quickly, even if the cut stems are placed in water. This is because the normal water transport system is poorly developed. As might be expected, there are also no stomata (breathing pores) on the leaves. Roots, which normally play a very important role in the absorption of nutrients and water from the substrate, are often also reduced and their main function is anchorage. The root hairs which function in absorption are often absent and roots themselves may be entirely dispensed with (e.g. Bladderwort). Many species have very specialised leaf shapes. The submerged leaves are often highly dissected or divided. This has the advantage of creating a very large surface area for absorption and photosynthesis. It also minimises water resistance and hence potential damage to the leaves. Heterophylly, where leaves of different shapes are produced depending on where on the plant they are, is common. This can create great problems for identification! In many cases, the submerged leaves are totally different to floating or emergent leaves on the same plant. The emergent leaves are usually much less divided, if not entire and have a more similar internal structure to those of land plants. Air-filled cavities often extend throughout the leaves and stems of aquatic plants, providing

an internal atmosphere. Certain aquatic fly and beetle larvae have a novel adaptation which allows them to take unusual advantage of this. A sharp appendage on the end of their abdomen is used to pierce into submerged plants, giving them access to the internal airfilled cavities as their own personal oxygen source.

Offwell Wetland Open Water Species List

Floating Plants

Fringed Water-lily Floating plants are of two types: those which are rooted with floating leaves (e.g. Water Lily) and those which are not rooted in the sediment, but just float on the surface (e.g. Duckweed). Floating leaves are generally tough because they have to withstand the weather and water movement. The green pigment-containing chloroplasts important for photosynthesis are restricted to the upper surface of the leaves which are the only surface to be well lit. Stomata (breathing pores), through which gas exchange takes place in the leaf, are also found only on the upper surface of the leaf. This upper surface often has a thick waxy cuticle to repel water and help to keep the stomata open and clear. Air-filled internal cavities are also often present. Terrestrial plants such as trees have to develop an enormous quantity of structural material in order to rise above all the other plants and collect the lion's share of the light available. Water lilies provide a neat example of a plant which has managed to do exactly the same thing, but with the minimum of structural material. Weak stems produce a massive floating canopy of leaves which dominate the local aquatic plant community just as effectively as trees dominate in a woodland. The difference lies in their external medium. Water provides all the necessary support, whereas air does not.

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A halophyte is a plant that naturally grows where it is affected by salinity in the root area or by salt spray, such as in saline semi-deserts, mangrove swamps, marshes and sloughs, and seashores. An example of a halophyte is the salt marsh grass Spartina alterniflora (smooth cordgrass). Relatively few plant species are halophytes - perhaps only 2% of all plant species. The large majority of plant species are "glycophytes", and are damaged fairly easily by salinity.[1] One quantitative measure of salt tolerance is the "total dissolved solids" in irrigation water that a plant can tolerate. Sea water typically contains 40 grams per litre (g/l) of dissolved salts (mostly sodium chloride). Beans and rice can tolerate about 1-3 g/l, and are considered glycophytes (as are most crop plants). At the other extreme, Salicornia bigalovii (dwarf glasswort) grows well at 70 g/l of dissolved solids, and is a promising halophyte for use as a crop. [2] Plants such as barley (Hordeum vulgare) and the date palm (Phoenix dactylifera) can tolerate about 5 g/l, and can be considered as marginal halophytes.[1] Adaptation to saline environments by halophytes may take the form of salt tolerance (see halotolerance) or salt avoidance. Plants that avoid the effects of high salt even though they live in a saline environment may be referred to as facultative halophytes rather than 'true', or obligatory, halophytes. For example, a short-lived plant species that completes its reproductive life cycle during periods (such as a rainy season) when the salt concentration is low would be avoiding salt rather than tolerating it. Or a plant species may maintain a 'normal' internal salt concentration by excreting excess salts through its leaves or by concentrating salts in leaves that later die and drop off.

Biology
Of the recognized 110 mangrove species, only about 54 species in 20 genera from 16 families constitute the "true mangroves", species that occur almost exclusively in mangrove habitats.[3] Demonstrating convergent evolution, many of these species found similar solutions to the tropical conditions of variable salinity, tidal range (inundation), anaerobic soils and intense sunlight. Plant biodiversity is generally low in a given mangal.[1] This is especially true in higher latitudes and in the Americas. The greatest biodiversity occurs in the mangal of New Guinea, Indonesia and Malaysia.[12]

[edit] Adaptations to low oxygen

A red mangrove, Rhizophora mangle

Above and below water view at the edge of the mangal

Red mangroves, which can survive in the most inundated areas, prop themselves above the water level with stilt roots and can then absorb air through pores in their bark (lenticels). Black mangroves live on higher ground and make many pneumatophores (specialised rootlike structures which stick up out of the soil like straws for breathing) which are also covered in lenticels. These "breathing tubes" typically reach heights of up to thirty centimeters, and in some species, over three meters. There are four types of pneumatophorestilt or prop type, snorkel or peg type, knee type, and ribbon or plank type. Knee and ribbon types may be combined with buttress roots at the base of the tree. The roots also contain wide aerenchyma to facilitate oxygen transport within the plant.

Salt crystals formed on grey mangrove leaf

[edit] Limiting salt intake

Red mangroves exclude salt by having significantly impermeable roots which are highly suberised, acting as an ultra-filtration mechanism to exclude sodium salts from the rest of the plant. Analysis of water inside mangroves has shown that 90% to 97% of salt has been excluded at the roots. Salt which does accumulate in the shoot concentrates in old leaves which the plant then sheds. Red mangroves can also store salt in cell vacuoles. White (or grey) mangroves can secrete salts directly; they have two salt glands at each leaf base (hence their namethey are covered in white salt crystals).

[edit] Limiting water loss

Because of the limited freshwater availability in salty intertidal soils, mangroves limit the amount of water that they lose through their leaves. They can restrict the opening of their stomata (pores on the leaf surfaces, which exchange carbon dioxide gas and water vapour during photosynthesis). They also vary the orientation of their leaves to avoid the harsh midday sun and so reduce evaporation from the leaves. Anthony Calfo, a noted aquarium author, observed anecdotally that a red mangrove in captivity only grows if its leaves are misted with fresh water several times a week, simulating the frequent tropical rainstorms.[13]

[edit] Nutrient uptake

The biggest problem that mangroves face is nutrient uptake. Because the soil is perpetually waterlogged, there is little free oxygen. Anaerobic bacteria liberate nitrogen gas, soluble iron, inorganic phosphates, sulfides, and methane, which makes the soil much less nutritious and contributes to mangroves' pungent odor. Prop root systems allow mangroves to absorb gases

directly from the atmosphere, and other nutrients such as iron, from the inhospitable soil. Mangroves store gases directly inside the roots, processing them even when the roots are submerged during high tide.

[edit] Increasing survival of offspring

Red mangrove seeds germinate while still on the parent tree

In this harsh environment, mangroves have evolved a special mechanism to help their offspring survive. Mangrove seeds are buoyant and therefore suited to water dispersal. Unlike most plants, whose seeds germinate in soil, many mangroves (e.g. Red Mangrove) are viviparous, whose seeds germinate while still attached to the parent tree. Once germinated, the seedling grows either within the fruit (e.g. Aegialitis, Acanthus, Avicennia and Aegiceras), or out through the fruit (e.g. Rhizophora, Ceriops, Bruguiera and Nypa) to form a propagule (a ready-to-go seedling) which can produce its own food via photosynthesis. The mature propagule then drops into the water which can transport it great distances. Propagules can survive desiccation and remain dormant for over a year before arriving in a suitable environment. Once a propagule is ready to root, its density changes so that the elongated shape now floats vertically rather than horizontally. In this position, it is more likely to lodge in the mud and root. If it does not root, it can alter its density and drift again in search of more favorable conditions.
hydathode (Gr. ydato: to make watery; odos: road, way) n. An epidermal structure, a pore, specialized for secretion, or for exudation, of water, usually from a leaf. Like stomata, hydathodes are surrounded by two crescent-shaped cells but these, unlike guard cells, do not regulate the size of the aperture. Hydathodes are used by the plant to secrete water under conditions in which transpiration is inhibited; for example, when the atmosphere is very humid. This process of water loss is called guttation

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Aerenchyma
From Wikipedia, the free encyclopedia Jump to: navigation, search

Aerenchyma is an air channel in the roots of some plants, which allows exchange of gases between the shoot and the root. The channel of large air-filled cavities provides a lowresistance internal pathway for the exchange of gases such as oxygen and ethylene between the plant above the water and the submerged tissues. Aerenchyma form in roots subject to anoxia such as occurs during flooding of plants and soil [1] . For example, Blom et al. (1994) investigated the adaptive responses of plants to flooding along the banks of the Rhine River, which included such morphological changes such as aerenchyma formation.

[edit] Aerenchyma formation

In maize, an aerenchyma is formed from highly selective cell death and dissolution in the root cortex during anoxia in the roots [2]. When plant roots are submerged or the surrounding soil flooded, hypoxia develops as soil microorganisms consume oxygen faster than diffusion occurs. Nitrification is inhibited as low oxygen occurs and toxic compounds are formed as anaerobic bacteria use nitrate, manganese, and sulfate as alternative electron acceptors [3]. The reduction-oxidation potential of the rhizhosphere decreases and metal ions such as iron and manganese precipitate [4]. Low oxygen generally stimulates trees and plants to produce ethylene [5]. Yet Visser et al.,1997 found ethylene slows down primary and adventitious root elongation and formation. Thus, in addition to supplying root tissues with oxygen, aerenchymas also assist in diffusing the accumulation of ethylene in order to prevent elongation inhibition (Visser et al. 1997).

[edit] How are aerenchyma formed?

Aerenchymas are formed by cell differentiation and collapse (lysigenous aerenchyma) or by cell separation without collapse (schizogenous aerenchyma). The differentiation or separation forms large continuous air spaces that allow diffusion of oxygen from shoot to root [6]. Different experiments defined how cell collapse occurs. Cell death was blocked by antagonists of phospholipid metabolism, of cytolsolic Ca2+ or Ca-calmodulin, and of protein kinases. By contrast, reagents that activate G-proteins, raise cytolsolic Ca2+ or inhibit phosphatases promoted cell death (two references He et al. 1996). An enzyme that was linked to this process was cellulase that assists in cell wall breakage. In maize a protein was found which is homologous to the enzyme XET, a protein that breaks the -1,4 links between glucans and xyulosyl the cross-linking molecule in plant cell walls [7].

[edit] What are some advantages of aerenchyma?

The large air-filled cavities provide a low-resistance internal pathway for the exchange of gases between the plant organs above the water and the submerged tissues. Some of the oxygen transported through the aerenchyma leaks through root pores into the surrounding soil. The resulting small rhizosphere of oxygenated soil around individual roots support microorganisms that prevent the influx of potentially toxic soil components [8] such as sulfide, iron, and manganese. Nitrifying bacteria provide the roots with a favourable nitrogen source [9] . During drought, aerenchymas allow plant roots to grow deeper for water, even through compacted layers; thick and tough roots are formed. As the roots dieback and decay, the resulting voids are paths in which new roots can grow and elongate when resources are available.

[edit] Any disadvantages?

Not all plants are able to develop aerenchymas. Aerenchymous roots may experience the following problems

Water and nutrient uptake may be less efficient; large intercellular spaces decrease the tissue available to transport water and nutrients from the root surface to the root xylem (Visser et al. 1996, 2000a). Large root diameters reduce biomass-to-surface ratio, resulting in less uptake of water and nutrients and the reduced opportunity to explore all microzones for nutrients. Some roots with aerenchymas are less likely to resist the physical strain of compacted soils. Those roots that penetrate and survive dense and compact drained soils have a higher bulk density and a strongly lignified layer of cells surrounding the aerenchyma, which strengthens the root. This dense, lignified layer prevents radial leakage of oxygen from the aerenchyma and may block some water and nutrient uptake (Colmer et al. 1998; Visser et al. 2000). During drought, roots with aerenchyma may be less tolerant to water stress as the open structure of the cortex is probably a low-resistance pathway for water vapor, as well as for air, thereby increasing the susceptibility of the root to water loss.