Unit 8: Sapwood and Heartwood

As a tree matures, the older xylem ceases to contain any living ray or axial parenchyma cells so any accumulating materials are converted into various extractive substances, including tannins, oils, gums, and resins. This centrally located wood is generally darker in color and is called heartwood. The heartwood is surrounded by lighter-colored sapwood containing living cells and starch. The wood of young trees is generally composed entirely of sapwood. As the tree ages, all the living cells in the central core of the trunk and large branches die, darken, and lose their physiological function. The most significant change in the transformation of sapwood into heartwood is death of ray and axial parenchyma cells. This can be observed in the gradual disappearance of nuclei in the parenchyma. Heartwood formation is also accompanied by anatomical changes such as an increased aspiration of pits in gymnosperm tracheids and the formation of tyloses in dicot vessel elements. As a general rule, heartwood is more resistant to penetration by preservatives than is sapwood. It follows, therefore, that heartwood is not suitable for timbers that will be impregnated with these fluids. Heartwood also is more difficult to season without checking. The primary reason for the drastic loss of permeability when going from sapwood to heartwood appears to be the irreversible aspiration of bordered pits and the encrustation and occlusion of the conducting elements by secondary metabolites and deposits. Heartwood cells and pit cavities become filled with hardened deposits, and the pit membrane tori often cover the bordered openings into the pit and cell cavities.

Unit 9: Periderm
In most woody perennial dicotyledons and gymnosperms, the original outer primary epidermal and cuticular covering is replaced in one year so that older stems and roots have an outer layer of secondary periderm. The periderm is usually initiated just after organ elongation is completed, although in some taxa it is much delayed. The periderm is not synonymous with the term bark. Bark is a nontechnical term that refers to all tissues exterior to the vascular cambium, therefore including secondary phloem. Bark refers to a number of tissue types of different origin, such as periderm and secondary phloem. In most woody plants, the vascular cambium is removed if the bark is stripped off. Bark formed during the early stages of the season's growth is often termed early bark or soft bark and consists mainly of sieve tubes and parenchymatous and suberized cells, but it does not include fibers or other strengthening cells. Toward the end of the growing season, late bark or hard bark is formed. It typically contains fibers and other sclerenchymatous cells, along with bark parenchyma and fewer and smaller sieve elements.

 Variation may be evident in the degree of development of periderm around the circumference of a single stem, which in extreme cases results in a " winged cork" condition. The mature periderm serves to protect the axis from attack by animals, fungi, and other pathogens. It also prevents the plant from drying out and insulates it from fire. The very thick (up to 1 if) fibrous periderm of the redwood tree (Sequoia) is a particularly effective fire shield because it lacks combustible resins and thus does not easily burn. Among different species, periderm morphology shows many differences in texture, color and thickness. Structurally the periderm consists of three components:o The initiating layer of the periderm is the phellogen, or cork cambium. o The cells produced to the outside of the phellogen are nonliving cork cells, technically called phellem. o In some plants, the phellogen forms one or more layers of internal tissue composed of living, cortexlike parenchyma cells referred to as phelloderm. The phellogen is a lateral meristem that forms at varying depths from the dedifferentiation of living, completely mature cells that resume division and become converted into a cambium. Depending upon the taxon, the phellogen may arise in the divisions of epidermal, cortical, or phloem parenchyma cellsof the stem. In the grapevine (Vitis), it originates as a result of cell divisions in the parenchyma of the metaphloem. In the vast majority of dicotyledons, the phellogen has its origin in subepidermal cortical layers, often in a discontinuous manner. This is referred to as a superficial origin of the cork cambium, as compared with a deep-seated origin within the inner cortex or secondary phloem. The root phellogen, in contrast, is always formed internally within the proliferating pericycle. Unlike the vascular cambium, the phellogen contains initials of only one type. These are more or less isodiametric in shape, and in transverse section appear radially flattened. As noted earlier, periclinal divisions of the phellogen produce two types of derivative tissue: the protective phellem (or cork) toward the outside and the phelloderm (or green cork) toward the inside. As a general rule, more radial files of cork cells are formed centrifugally than phelloderm cells are formed centripetally. Collectively, the three layers constitute the periderm. With formation of the periderm, the primary epidermis becomes effectively isolated from internal tissues. The epidermis cracks and is sloughed off, although the dead and collapsed epidermis may cling to the periderm for some time. Mature cork cells are usually arranged in compact, radial rows and are nonliving. They have an air space in the center that contributes to the buoyancy of cork in water. The air spaces also push compressed cork back to its original shape. Cell walls are thin or thick and without pitting and characteristically are full of a waxlike substance called suberin. The deposition of suberin typically occurs in layers alternating with other waxes, resulting in alamellate structure of suberized walls when viewed in thin sections. In addition to suberin, all barks contain cellulose, hemicellulose, lignin, darkstaining polyphenols, and other extractives. Suberin and lignin are the principal components of cork, contributing about 40% and 22%, respectively, of its dry weight. The presence of

large quantities of suberin in the walls of cork cells makes them poor conductors of heat and highly impervious to gases and liquids. These qualities make cork an effective barrier against the outside environment; thus cork is sometimes utilized in the manufacture of insulation board. The internal derivatives of the phellogen rarely comprise more than a few loosely organized rows that resemble cortical cells. In contrast to phellem, mature parenchymatous phelloderm cells retain their protoplasts with nuclei. The fact that these cells may have chloroplasts accounts for their designation as green cork. Phelloderm is not present in all woody plants. To exchange gases and water, woody plants almost universally develop small structures known as lenticels within the periderm. Lenticels form on stems and even some fruits. Lenticels commonly arise beneath stomata or between stoma, usually just prior to or at the same time as the first periderm. As the lenticel develops, the epidermis is ruptured, and the mass of lenticel tissue is exposed. The inner boundary of the lenticel is a meristematic zone that is continuous with the phellogen. A fully developed lenticel is distinguished from the phellem by having a mass of loosely arranged, thin-walled, and unsuberized cells with abundant intercellular spaces. This tissue is called complementary tissue. At maturity the cells making up the complementary tissue lose their protoplasts and become colorless. Complementary tissue can occur as a somewhat uniform and compact tissue, or they can be rather loosely organized. In some plants (e.g., Ulmus, Alnus, Betula, and Sorbus), the complementary tissue is horizontally traversed by bands of densely arranged, suberized cells called closing layers. Beneath the lenticel, the cortical air spaces become continuous with those in the complementary region. As axis diameter increases as a result of the production of secondary tissues, the periderm in most species is subjected to stresses that cause it to crackand split. Anticlinal divisions within the phellogen enable the cambium to increase in girth in concert with the increasing width os the vascular cylinder. As the axis grows, additional successive periderms can arise at any time interior to the one most recently produced. As this occurs, new segments of cork cambium develop in progressively older regions of the axis, often developing in overlapping layers. The isolated older tissues that are formed by the development of deeper and deeper cork cambia are called rhytidome. These tissues may include periderm and masses of cortical and phloem cells that have been successively cut off. Very little is known about the seasonal pattern of activity of the cork cambium, what stimulates the cambial cells to divide, and what controls the pattern of phellogen development. Bark surface features aredetermined by the developmental pattern of the cork cambium. In some trees, the bark is thin (0.5 in.), continuous, and smooth, and the original phellogen remains functional over an extended period. In these cases, the phellogen increases in girth by frequent anticlinal divisions.

When trees develop successively more internal meristems, however, the outer bark becomes irregularly split and exfoliates in large sheets or in small, irregular segments. Trees in which phellogens are renewed in concentric, continuous layers shed their barkin thin strips, papery layers or as cylinders. This type of bark pattern is called ring bark. Replacement phellogens that form in overlapping layers produce a scale bark surface pattern. Economic importanceThe most commercially valuable cork is derived from the outer bark of the cork oak (Quercus suber), an evergreen tree from the Mediterranean region. A uniform, high quality cork layer forms over a period of several years and is periodically stripped from the tree. If the harvesting is done carefully, the inner phellogen remains undamaged, and a layer of new growth cork is formed. Everyone is familiar with the most common use of cork, namely cork bottle stoppers. The anatomical characteristics of cork make it ideal for this purpose. Cork possesses lenticels, however, that extend through the tissue allowing for gas exchange between the outside and inner tissues. For this reason, bottle stoppers must be cut out at right angles to the surface of a piece of cork so that the lenticels extend horizontally rather than vertically, which would allow leakage of the contents. Barks also are used as a source of valuable fibers, resins, tannins, latex, medicines, poisons, flavors, and hallucinogenic substances. The remarkable properties of Prunus serrula bark has recently been described. The thin, tough, flexible, and semitransparent bark of this species has been compared to a strong natural polymer film. Unlike most barks that have cuboidal or slightly expanded cork cells among the tangential axis, the cork cells of Prunus serrula have considerable tangential enlargement many times the radial dimension. The bark also shows exceptionally high toughness values and high density.

Unit 10: Ecological Anatomy

Xeromorphic Leaves In addition to physiological differences, xerophytic plants typically possess one or more striking morphological and anatomical modifications to water stress, many of these are associated with leaf structure. Xeromorphic foliar structure is sometimes correlated with the absence of certain nutrients in the soil. Low soil nitrogen levels can result in increased lignification of the leaf epidermis and nonveinal sclerenchyma. High nitrogen levels can cause a decrease in the amount of wood formed. Plants that live in habitats where the supply of available water is deficient have been classified into broadly defined categories on the basis of the different adaptive strategies they have developed:

1. Drought-escaping species are plants with a compressed growth cycle that complete vegetative growth and reproduction within the short period of time when conditions are favorable. 2. Drought-evading species are plants that are able to reduce water loss, or compensate for water loss, by possessing some specialized structural feature or set of features, such as an extensive root system. 3. Drought-enduring species are plants that are able to survive even when water uptake cannot take place or is severely reduced. This group includes species with a number of highly divergent specializations such as temporary leaf loss, changes in leaf angle and the rolling or folding of leaf blades, or permanent adaptations such as succulence or leaves reduced to spines. The leaves of some xerophytic grasses contain enlarged epidermal cells that store water and occur in dispersed longitudinal rows. These cells are called bulliform cells and under conditions of water deficit they lose turgor and thus constrict in upon themselves, causing the grass lamina to fold or roll inward edge to edge. Bulliform cells may occur only on the upper surface or both upper and lower surfaces. As a result, the entire upper leaf surface forms a protective barrier to air movement that limits water loss. Bulliform cells are recognizable by the cell wall ridges that are present when the cells are constricted. The effects of environmental factors on leaf morphology and anatomy have produced a wide variety of adaptations in different taxa. Some features that probably contribute to a reduction in the rate of loss of absorbed water are:i. reduction in leaf size, ii. increased thickness of the outer walls of epidermal cells, iii. increased thickness of cuticle, iv. increased trichome density, v. reduction in stomatal pore area, and vi. stomata recession below the blade surface (or confined to pits or grooves on the undersurface of the blade), Other characteristics that probably help the plant to tolerate dehydration and deal with water stress are:o increase in mechanical cells and wall lignification, o increased succulence and water storage capacity, and o the accumulation of mucilage,. Halophytic Leaves Halophytic plants, or halophytes, are those species able to tolerate saline environments. This adaptation is sometimes considered a special type of xerophytism because of the nature of the environments they inhabit. Halophytic species often show a diversity of structural and physiological adaptations that include succulent leaves with specialized "colorless" storage cells or "window cells" that increase the plants capacity for salt accumulation.

 Some species have secretory cells or tissues, enlarged tracheids terminating vein endings, structures such as salt glands and salt hairs, and a hypodermis. Halophytic plants concentrate salt in their tissues, moving it from the soil into and through the plant. The leaves of the salt-tolerant maritime mangrove (e.g., Avicennia germinans) contain large, tightly packed, multiple rows of specialized hypodermal cells that play a probable role in salt accumulation and storage. Because salt cannot be allowed to accumulate without limit, many halophytes (including mangroves) have special surface glands and hairs to remove NaCl from the underlying mesophyll cells and then actively secrete it on the leaf surface. The salt glands of Avicennia are formed in shallow pits on the upper leaf surface, whereas on the lower surface they are not sunken. Individual glands consist of two to four highly vacuolated basal cells, a cutinized stalk cell, and a minimum of eight terminal cells that are covered by a thin, perforated cuticle that is separated from the cell wall apically. All gland cells are interconnected by plasmodesmata. Salt is deposited in the subcuticular cavity of the gland and then ultimately reaches the leaf surface through the cuticular pores. The structurally complex salt glands of the facultative halophyte Limonium gmelini (Plumbaginaceae) are composed of 16 glandular cells overlying 4 collecting cells that are connected to mesophyll cells by plasmodesmata. A layer of cutin surrounds the collecting cells except at pore interruptions. Other halophytes such as Atriplex (Chenopodiaceae) have salt hairs on the surface. These specialized hairs are composed of an enlarged terminal bladder cell that contains a large saltaccumulating and salt-storing central vacuole. The bladder cell is attached to the leaf epidermis by a stalk cell, through which the salt is actively transported. Eventually these salt-containing cells burst and crystalline salt is deposited on the epidermal surface.

Function: o The surface salt provides a reflective coating that shades the leaf from direct sunlight and also decreases the palatability of the plant to herbivores. o The sharp salt crystals also may function as a mechanical deterrent to herbivory. In the halophyte Aster tripolium, stomata close in response to high Na+ concentrations. This response prevents excessive accumulation of Na+ within the shoot by controlling transpiration rates. Hydromorphic Leaves Water plants show all stages of this transition with emergent, floating, or permanently submerged leaves. All these hydrophytic leaf types vary considerably in response to changes in habitat, although all are characterized by marked increases in aerenchyma, parenchyma tissue with particularly large intercellular spaces.

 Whereas emergent upright aquatic leaves are otherwise similar in structure to those of nonaquatics, floating and submerged leaves possess a number of distinctive features that serve such functions as gas exchange and flotation. Free-floating plants such as the duckweed (Lemna) possess leaves modified to trap air and prevent waterlogging by having large, specialized trichomes and a thick cuticle. The structural characteristics in aquatic plants are mainly related to a reduction of the protecting, supporting, and conducting tissues and the presence of air chambers in tissues of the leaf, stem, and root. Epidermal cells typically possess chloroplasts that increase light absorption and photosynthesis. Anatomical adaptations represent responses to excessive water content, which implies a decreased oxygen supply. The anatomy of leaves of the pondweed Potamogeton illustrates this syndrome; the leaves possess weakly developed supporting and conducting tissues and prominent intercellular air spaces. The stems of submerged aquatics like Potamogeton also contain large gas lacunae that provide pathways for the transport of oxygen to the roots. Although such pathways are continuous down the stem, they are interrupted by thin, perforated diaphragms. Experimental work has shown that these structures act to control leaks and provide most of the resistance in the mass flow of gases but contribute little resistance to diffusive gas transport. With the notable exception of the genus Brasenia (Cabombaceae), likely stages in tissue and cell reduction have not been well documented among aquatic plants. Cuticle on the epidermis is generally absent or is extremely thin. The epidermis is not primarily protective in this case but is composed of epidermal cells that are modified into transfer cells in such a way that they facilitate the absorption of gases and nutrients directly from the water. Chloroplasts occur in the epidermal cells, although they decrease in abundance with increasing water depth and with decreasing light. Large air passages filled with gases are common in hydrophytes of this type. In addition to providing very efficient mechanical stabilization to organs, the oxygen given off in photosynthesis as well as that from the air is stored and transported in these spaces, to be used again in respiration. The carbon dioxide from respiration also is held and used in photosynthesis. The function of air chambers in buoying some floating leaves is equally clear. Sclerenchyma is generally reduced or absent, although sclerenchymatous cells are conspicuous in the leaves of some marine angiosperms. Stomata are completely lacking in permanently submerged leaves. In floating leaves the lamina has retained a dorsiventral structure, and functional stomata are restricted to the adaxial (upper) surface of the lamina. A number of unrelated aquatic plants develop leaves of two distinct morphologies and anatomies on the same stem--one form of leaf submerged and the other form aerial. This condition is known as heterophylly or leaf dimorphism. Shoot apices of submerged and aerial axes appear similar. Leaves originating from an aerial shoot apex are usually undissected, with an entire, lobed, or toothed blade, with stomata, and with a biracial mesophyll with a palisade zone. Leaves arising from submerged apices are highly dissected, are essentially devoid of stomata and cuticle, have an undifferentiated mesophyll that often lacks palisade cells, and have reduced venation patterns. Among species with whorled phyllotaxy, the aerial stems typically have fewer leaves in each whorl as compared with the

submerged segment of stem. In these cases, increase in the number of dissected leaves in a whorl increases total leaf surface area and the total assimilating area. Experiments with the mermaid weed Proserpinaca palustris have demonstrated that conditions of the photoperiod and other environmental factors can influence the form of leaf produced at the apex. We know from experiment that the developmental determination of heterphyllous leaf types involves the action of the hormone abscisic acid (ABA). When exposed to ABA, pondweeds form floating or aerial leaves instead of submerged leaves. Determination in these cases is a gradual process that is not finalized until late in development, when the leaf is almost fully expanded. Growing leaf primordia placed in different environmental regimes (aerial and submerged) at different stages of development will become either entirely one leaf type or the other, or they may show intermediate features, depending upon the stage of leaf development at which the shift in environment was made. If the transfer is made at an early stage, the leaf will resemble the leaf type characteristic of the new environment. If the shift is made later in development, the leaf will display intermediate features. TABLE: Common Anatomical Differences Between Leaves of Hydrophytes and Xerophytes

Roots
Xeromorphic Roots The root systems of certain xerophytes are widely spreading and shallow for optimal water absorption. A short tuberized root is common in some xeric taxa and there is an absence of root hairs under drought conditions. Xeromorphic roots also can be succulent because they contain large water storage cells in the cortex. Some plants from dry regions produce roots that develop a thick, sclerified, and exfoliating bark that cracks and splits off in large sheets. Cactaceae and other succulents conserve water during prolonged dry periods by having few, if any, lateral branch roots; the entire root system may be covered by layers of thick bark. As the soil water potential increases, the dormant root system quickly forms new lateral roots, termed rain-induced roots, that are able to absorb large quantities of water. Some plants that grow in nutrient-poor soils form dense clusters of determinate lateral rootlets. These are sometimes called proteoid roots, after their occurrence in some Proteaceae where they are believed to be involved in phosphate acquisition. Specialized cluster roots also occur in members of the Casuarinaceae, Myricaceae, Betulaceae, and Moraceae. Hydromorphic Roots The roots of aquatic plants range from well developed like those of terrestrial species through various degrees of specialization, including root dimorphism with respect to the presence or absence of root hairs. Because mechanical support and conducting efficiency are no longer necessary, vascular systems are to be ontogenetically and phylogenetically reduced and narrow, with weakly lignified tracheary elements. This reduction in vascular tissue tends to be correlated with the degree of specialization of the plant. Emergent aquatics possess the least reduction in the quantity of conducting tissues, whereas submerged plants have the most reduced. In cases of extreme reduction, the stele can be only 50 lam in diameter and composed of very few sieve tube elements and tracheary elements that are thin-walled and probably vestigial. I n hydromorphic roots, the sieve tube elements are commonly distributed in an unusual pericyclic manner. Prominent cortical air spaces are formed by the breakdown or separation of alternate plates of cells and are continuous with those in the stem and leaves. In plants with a well-developed lacunose ground tissue, there is a ready diffusion of oxygen from the stems and leaves down to the roots that are embedded in an oxygen-deficient substrate. Root growth, therefore, is not normally oxygen limited.

 In rare instances, carbon dioxide is known to be absorbed by the roots and the gas moves through the air spaces to the leaves where it is photosynthetically fixed. Aquatic plants lacking abundant air space systems in their vegetative organs normally grow in rapidly moving streams and near waterfalls where oxygen appears not to be a limiting factor. The Podostemaceae are one of these families. The air chambers in aquatic plants are regularly traversed by septa or diaphragms that are solid or porous and sometimes constructed of stellate parenchyma. Diaphragms are thought to serve three functions: i. to provide strength and stabilization to organs and prevent collapse of the air chambers, ii. to restrict the movement of water when the organ is damaged and internal flooding occurs, and iii. to provide a lateral transport pathway across the cortex.